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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


us  ISSN  0027.4100 


bulletin  OF  THE 

Museum   of 

Comparative 
Zoology 


A  Review  of  the  North  American 
Fossil  Amiid  Fishes 


JOHN  R.  BORESKE,  JR. 


HARVARD  UNIVERSITY 

CAMBRIDGE,  AAASSACHUSEHS,  U.S.A. 


VOLUME  146,  NUMBER  1 
18  JANUARY  1974 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED  BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 

Bulletin  1863- 

Breviora  1952- 

Memoibs  1864-1938 

JoHNSONiA,  Department  of  Molliisks,  1941- 

OccAsioNAL  Papers  on  Mollusks,  1945- 


Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.     Fishes  of  tlie  Gulf  of  Maine. 
Reprint. 

Bnies,  C.  T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.     Classification  of 
Insects. 

Creighton,  W.  S.,  1950.    The  Ants  of  North  America.     Reprint. 

Lyman,  C.  P.,  and  A.  R.  Dawe  (eds.),  1960.    Symposium  on  Natural  Mam- 
malian Hibernation. 

Peters'. Check-list  of  Birds  of  the  World,  vols.  2-7,  9,  10,  12-15. 

Sprinkle,  J.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echinoderms. 

Turner,  R.  D.,  1966.    A  Survey  and  Illustrated  Catalogue  of  the  Teredinidae 
(Mollusca:  Bivalvia). 

Whittington,  H.  B.,  and  W.  D.  I.  Rolfe  (eds.),  1963.    Phylogeny  and  Evolu- 
tion of  Crustacea. 

Proceedings  of  the  New  England  Zoological  Club  1899-1948.     (Complete 
sets  only.) 

Publications  of  the  Boston  Society  of  Natural  Histor>'. 


Authors  preparing  manuscripts  for  the  Bulletin  of  the  Museum  of  Comparative 
Zoology  or  Breviora  should  send  for  the  current  Information  and  Instruction  Sheet, 
available  from  Editor,  Publications  OflBce,  Museum  of  Comparative  Zoology, 
Harvaid  University,  Cambridge,  Massachusetts  02138,  U.S.A. 


(g)  The  President  and  Fellow*  of  Harvard  Colleee  1974 


A  REVIEW  OF  THE  NORTH  AMERICAN 
FOSSIL  AMIID  FISHES 

JOHN  R.  BORESKE,  JR.i 


CONTENTS 

Abstract   1 

Introduction  2 

Acknowledgments  2 

Abbreviations  3 

A7?Jia   calva   3 

Nomenclature    3 

Ecology    4 

Geographic   Distribution    4 

Pleistocene  Occurrences  4 

Diagnosis  5 

Morphometries    5 

Methods  6 

General  Proportions  and  Growth  8 

Comparisons  with  Fossil  Forms  10 

Discussion    17 

Meristics    18 

Supravertebral  Scale  Rows  18 

Branchiostegal    Rays    20 

Fin  Rays  20 

Vertebral  Elements  25 

Vertebral  Column  of  Amia  calva  28 

Vertebral  Features  28 

Vertebral  Dimensions  33 

Valid  North  American   Fossil   Genera   and 

Species  37 

Amia   fragosa    37 

Amia   uintaensis   47 

Amia  cf.  uintaensis  64 

Amia  scutata  66 

Amia  cf.  scutata  70 

Amia  cf.   calva  72 

Amiidae   incertae   sedis   72 

Specimens  Removed  from  the  Amiidae  74 

Summary  and  Conclusions  75 

Literature  Cited  81 

Plates   84 

Abstract.     North  American  amiid  fishes  range 

from  Cretaceous  ( Albian )  to  Recent.    Amiids  are 


1  Museum    of    Comparative    Zoology,     Harvard 
University,  Cambridge,  Massachusetts  02138 

Bull.  Mus.  Comp.  Zool,   146(1):   1-87,  January,   1974 


common  fossils  in  Late  Cretaceous  and  Tertiary 
freshwater  deposits  and  apparently  occupied  a 
habitat  much  like  that  of  the  Recent  species  Amia 
calva.  Morphometric,  meristic,  and  cranial  char- 
acters of  articulated  specimens  from  the  Fort 
Union  Fonnation  (Paleocene),  Green  River  For- 
mation (Eocene),  Florissant  Fonnation  (Oligo- 
cene).  Pawnee  Creek  Formation  (Miocene),  and 
a  Recent  A.  calva  sample  from  Wisconsin  have 
been  used  here  in  an  attempt  to  revise  the  taxon- 
omy and  evolutionary  history  of  the  group. 

Whereas  seven  genera  and  twenty-three  species 
of  fossil  amiids  have  been  described  on  the  basis 
of  disarticulated,  often  isolated  elements,  only 
three  taxa  have  heen  described  from  complete  or 
partially  complete  material.  Amia  fragosa  (Late 
Cretaceous  to  Middle  Eocene),  A.  uintaensis  (Pal- 
eocene to  Early  Oligocene),  A.  scutata  (Early  to 
Middle  Oligocene),  and  A.  calva  (Middle  Plio- 
cene to  Recent)  are  here  considered  the  only 
valid  taxa.  Amiid  remains  are  first  known  in  the 
North  American  fossil  record  from  the  Early  Cre- 
taceous (Albian)  Paluxy  Formation  of  Texas. 
This  disarticulated  material  shows  resemblances 
both  to  A»ii«  and  to  the  Late  Mesozoic  European 
genera  Uroclcs  and  Arniopsis.  Paramiatus  gurleyi 
(Romer  and  Fryxell,  1928)  from  the  Green  River 
Formation  of  Wyoming  is  a  synonym  of  A.  frag- 
osa. Tlie  differences  between  Amia  and  the  large 
Early  Cenozoic  form  Protamia  are  insufficient  for 
recognition  of  Protamia  as  a  genus  distinct  from 
Amia.  The  Eocene  and  Oligocene  forms  Protamia 
media,  Pappichthys  medius,  P.  plicatus,  P.  scler- 
ops,  P.  laevis,  P.  symphysis,  P.  corsonii,  Amia 
whiteavesiana,  and  A.  macrospondyla  are  s\monyins 
of  A.  uintaensis;  they  were  based  on  undiagnostic 
cranial  and  vertebral  characters.  Morphometric 
and  meristic  similarities  indicate  that  little  evi- 
dence exists  for  maintaining  separate  Oligocene 
species  A7nia  scutata  and  A.  dictyocephala.  Amia 
exilis  is  a  synonym  of  Amia  scutata;  it  was  based 
on  undiagnostic  vertebral  characters.  A.  scutata 
is  morphometrically  distinguishable  from  A.  calva 
only  on  the  basis  of  a  slightly  larger  head/stan- 

1 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


dard-length  ratio.  The  Eocene  ta.xa  Amia  de- 
pressus,  A.  newherrianus,  A.  gracilis,  and  Htjpamia 
elegans  are  nomina  diibia. 

Comparison  of  the  fossil  forms  with  the  Recent 
Amia  calva  suggests  the  following  ta.xonomic  and 
possible  phylogenetic  relationships:  ( 1 )  Amia  frag- 
osa  survived  until  the  Middle  or  Late  Eocene, 
with  no  phylogenetic  affinities  with  the  modern 
form;  (2)  Amia  iiintaensis  appears  to  be  closer 
than  Amia  fragosa  to  the  ancestry  of  Amia  calva, 
which  evolved  through  an  intermediate  fonn  such 
as  Amia  scutata;  (3)  establishment  of  the  Recent 
species  Amia  calva  had  begun  at  least  by  the  be- 
ginning of  the  Pliocene;  and  (4)  diere  are  simi- 
larities in  the  Paleocene  and  Eocene  amiid  fossil 
record  of  North  America  and  Europe. 

INTRODUCTION 

Aviia  is  a  genus  of  freshwater  fishes  that 
includes  one  of  two  extant  representatives 
of  the  holostean  level  of  organization.  It 
includes  a  number  of  species  of  which  only 
Amia  calva  exists  today;  other  forms  of 
Amia  are  found  in  the  fossil  record,  and 
extend  from  the  Late  Cretaceous  to  ap- 
proximately the  Middle  Pliocene.  This 
study  is  an  attempt  to  determine  the  taxon- 
omic  and  phylogenetic  relationships  among 
the  various  species  of  Amia.  It  is  established 
on  osteology  as  well  as  on  morphometric 
and  meristic  data  from  both  Recent  and 
fossil  forms.  This  data  is  used  to  compare 
the  available  features  of  the  fossil  forms 
with  Recent  Amia  calva  and  to  detennine 
the  validity  of  previous  descriptions  based 
on  various  osteological,  morphometric,  or 
meristic  character-states. 

Until  recently,  a  major  difficulty  in  inter- 
preting the  taxonomy  of  fossil  amiids  has 
been  the  paucity  of  articulated  specimens. 
Five  genera  and  twenty-one  species  of  fossil 
forms  have  been  described  from  disarticu- 
lated, often  isolated,  elements  (Table  19); 
only  two  taxa  have  been  described  from 
articulated  specimens:  Paramiatus  ii^iirleyi 
(Romer  and  Fryxell,  1928)  and  Amia 
scutata  (Osborn  et  al.,  1875).  Recent  works 
by  Estes  (1964)  and  Estes  and  Berberian 
( 1969 ) ,  based  on  disarticulated  elements 
from  the  Late  Cretaceous  Lance  and  Hell 
Creek  formations,  are  the  only  published 
studies  of  Amia  fragosa,  although  O'Brien 
(1969)    completed   an   M.A.   thesis  on  the 


osteology  of  A.  frap.osa,  describing  articu- 
lated specimens  from  the  Late  Cretaceous 
Edmonton  Formation  of  Alberta. 

Much  more  articulated  material  is  now 
available  and  provides  more  detailed  in- 
formation on  the  cranial  and  postcranial 
anatomy  of  amiids.  These  specimens  have 
been  useful  in  this  revision  of  the  taxonomy 
as  well  as  in  the  determination  of  possible 
relationships  to  European  and  Asian  forms. 

In  an  attempt  to  understand  the  evolution 
and  interrelationships  of  the  fossil  and  Re- 
cent amiids,  a  growth-series  study  has  been 
made  on  a  Recent  A.  calva  sample  from 
Wisconsin,  and  is  compared  moi"phometri- 
cally  and  meristically  with  the  fossil  forms. 
A  great  number  of  fossil  specimens,  includ- 
ing the  holotypes  and  paratypes  of  all  North 
American  amiid  species,  have  been  exam- 
ined. Several  European  taxa  have  been 
studied  at  the  British  Museum  ( Natural 
History),  London;  Museum  National  d'His- 
toire  Naturelle,  Paris;  and  the  Institut  Royal 
des  Sciences  de  Belgique,  Brussels. 

ACKNOWLEDGMENTS 

I  am  especially  grateful  to  Professor 
Richard  Estes  (University  of  California  at 
San  Diego)  for  his  advice  and  criticism  in 
the  preparation  of  this  manuscript.  Cecile 
Janot- Poplin  and  Sylvie  Wenz  ( Museum 
National  d'Histoire  Naturelle,  Paris),  and 
Karel  Liem  ( Museum  of  Comparative  Zo- 
ology) read  the  manuscript  and  offered 
criticisms  that  substantially  improved  the 
text. 

Additional  thanks  are  due  to  Donald 
Baird  ( Princeton  University ) ,  Henry  Booke 
and  Bany  Cameron  (Boston  University), 
William  J.  Hlavin  (Cleveland  Museum  of 
Natural  History),  Farish  A.  Jenkins,  Jr. 
(Museum  of  Comparative  Zoology), 
Charles  Meehan  ( Chamberlayne  College), 
Robert  R.  Miller  (University  of  Michigan), 
David  Pariis  ( New  Jersey  State  Muse- 
um), Colin  Patterson  (British  Museum  of 
Natural  History),  Clayton  Ray  (National 
Museum  of  Natural  History),  Bobb  Schaef- 
fer  (American  Musevun  of  Natural  History), 
Hans-Peter    Schultze     ( Geologisch-Paleon- 


I 


Fossil  Amiids  •  Borcske 


tologisches  Institiit  der  Gcorg-Aiigust-Uni- 
vcrsitiit,  Gottingen),  Keith  Thomson  (Yale 
University),  and  Hainer  Zangerl  (Field 
Museum  of  Natural  History)  for  their  help- 
ful suggestions.  I  am  also  grateful  to  Leslie 
Whone  for  preparation  of  tables,  and  to  Siri 
Falck-Pedersen  Boreske,  Laszlo  Meszoly, 
and  Charles  Chamberlain  for  illustrations. 
This  study  was  supported  by  grants  from 
Sigma  Xi,  Marsh  Fund,  and  the  Albion 
Foundation. 

ABBREVIATIONS 

AMNH — American  Museum  of  Natural 
History,  New  York,  New  York. 
ANSl^ — Academy  of  Natural  Sciences  of 
Philadelphia,  Philadelphia,  Pennsylvania. 
BMNH — British  Museum  (Natural 
History),  London,  England. 
CM — Carnegie  Museum,  Pittsburgh, 
Pennsylvania. 

F:  AM — Frick- American  Museum 
Collection,  New  York,  New  York. 
FHKSM— Fort  Hays  Kansas  State  Museum, 
Hays,  Kansas. 

FMNH — Field  Museum  of  Natural  History, 
Chicago,  Illinois. 

FSM — Florida  State  Museum,  Gainesville, 
Florida. 

MCZ — Museum  of  Comparative  Zoology, 
Harvard  University,  Cambridge, 
Massachusetts. 

MNHN — Museum  National  d'Histoire 
Naturelle,  Paris,  France. 
NMC — National  Museum  of  Canada, 
Ottawa,  Canada. 

PU — Museum  of  Natural  History, 
Princeton   University,   Princeton,   New 
Jersey. 

ROM — Royal    Ontario    Museum,    Toronto, 
Canada. 

SMUSMP— Shuler  Museum  of  Paleontol- 
ogy, Southern  Methodist  University,  Dallas, 
Texas. 

UA — University  of  Alberta  Museum, 
Edmonton,  Canada. 
UCMP — Museum  of  Paleontology, 
University  of  California,  Berkeley, 
California. 


UMM — West  Texas  Museum,  University  of 

Texas,  El  Paso,  Texas. 

UMMP — l^ni\ersity  of  Michigan  Museum 

ot  Paleontology,  Ann  Arbor.  Michigan. 

UMMZ — University  of  Michigan   Museum 

of  Zoology,  Ann  Arbor,  Michigan. 

USNM— National  Museum  of  Natmal 

History,  Wa.shington,  D.C. 

YPM — I'eabody  Museum  of  Natural 

History.  Yale  University,  New  Haven, 

Connecticut. 

AMI  A  CALVA   LINNAEUS,   1766 

Amid  calvii  is  the  only  extant  species  of 
the  family  Amiidae.  It  is  a  predaceous  fish 
that  exclusively  inhabits  fresh  waters  of 
the  eastern  LTnited  States.  Except  for  the 
gar,  Lepisosteus,  Amia  calva  is  the  onl\' 
other  living  representative  of  the  holo.stean 
fishes.  Its  common  name,  "bowfin,"  refers 
to  the  long  dorsal  fin  that  arches  in  a  bow 
over  most  of  the  length  of  the  fish's  back. 
Amia  calva  has  previously  been  known  as 
the  dogfish,  marshfish,  mudfish,  grindle,  or 
lawyer. 

The  osteology  of  Amia  calva  has  been 
extensively  described  and  discussed  by 
Schufeldt  (18S5),  Bridge  (1S77),  Allis 
(1889,  1897),  and  Goodrich  (1930).  The 
following  discussion  is  limited  only  to  the 
nomenclatural  problems,  ecology,  geo- 
graphic distribution,  and  character-states 
of  Amia  calva  that  are  relevant  to  study  of 
the  fossil  forms. 

Nomenclature 

Jordan  and  Evermann  (1896)  noted  that 
although  Linnaeus  (1766)  had  applied  the 
binomial  name  Amia  calva  to  the  genus, 
Gronow  (1763)  had  earlier  used  Amia  as 
a  nonbinomial  name  for  fishes  presently 
classified  as  Apoiion  Lacepede.  They  fur- 
ther suggested  that  should  Gronow's  earlier 
ipplication  of  the  name  be  given  prece- 
dence and  transferred  to  Apof^on,  then 
Ainiatus  Rafinesque  (1815)  should  replace 
Amia  Linnaeus.  Jordon  (1906)  stated  that 
this  transfer  of  names  was  a  necessary  com- 
pliance with  the  rules  of  nomenclature,  but 
later    (1919),   although   citing  Opinion   20 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


( 1910 )  of  the  International  Commission  on 
Zoological  Nomenclature  which  favored 
Gronovv's  priority,  Jordan  found  tlie  trans- 
fer of  names  inconvenient,  for  most  autliors 
had  rejected  Gronow's  names  as  pre- 
Linnaean.  In  1925,  Jordan  recommended  to 
the  Commission  that  certain  names  of 
Gronow  supported  by  Opinion  20  be  re- 
jected in  favor  of  the  more  accepted 
Linnaean  terminology.  The  Commission's 
Opinion  89  ( 1925 )  resolved  ( among  others ) 
the  nomenclatural  problem  of  Amia,  by  con- 
curring with  Jordan's  recommendation  that 
".  .  .  Amia  Gronow  be  set  aside  in  favor  of 
Amia  Linnaeus,  even  if  other  names  of 
Gronow  are  allowed."  Rafinesque's  name 
Amiatus  is  then  a  junior  synonym  of  Aynia 
Linnaeus. 

Some  later  workers  seem  to  have  been 
unaware  of  Opinion  89.  Thus  Hussakof 
( 1932 )  accepted  the  validity  of  the  transfer 
of  the  name  A7nia  Gronow  to  the  percoid 
teleost  Apo^on.  Romer  and  Fryxell  (1928) 
named  their  fossil  amiid  from  the  Eocene 
Green  River  Formation  Paramiatus  instead 
of  Paramia,  and  Whitley  ( 1954 )  changed 
the  name  of  Lehman's  (1951)  fossil  amiid 
from  the  Eocene  of  Spitzbergen  from  Pseu- 
damia  to  Pseudamiatus.  The  latter  is  invalid 
as  Pseudamia  was  a  valid  name  in  itself  and 
Pseudamiatus  is  its  junior  synonym  regard- 
less of  the  Amia- Amiatus  controversy. 

Ecology 

Aside  from  notes  regarding  breeding,  diet, 
and  zoogeographical  occurrences,  little  has 
been  written  in  the  past  50  years  about  the 
ecology  of  Amia  calva.  Dean  (1898)  and 
Reighard  ( 1903 )  have  made  the  only  ex- 
tensive published  investigations  of  the 
habits  and  habitat  of  the  fish.  A  thorough 
study  of  the  biology  of  A.  calva  throughout 
its  range  is  long  overdue. 

Geographic  Distribution 

The  distributional  map  of  Amia  calva 
( Fig.  1 )  is  based  on  information  drawn 
from  Hubbs  and  Lagler  (1967),  and  Blair 
et  al.  ( 1968 ) ,  and  from  examinations  of 
unpublished  records  at  the  Ohio  State  Uni- 


versity Museum  of  Zoology,  Museum  of 
Comparative  Zoology,  and  the  University 
of  Michigan  Museum  of  Zoology.  The  dis- 
tribution limit  is  a  flexible  boundary  allow- 
ing for  seasonal  occurrences  and  other 
natural  variations.  The  known  northern 
limit  of  A.  calva  extends  from  the  Missis- 
sippi drainage  system  in  Minnesota  south  of 
Duluth,  eastward  through  Lake  Nipissing 
and  the  Ottawa  River  to  the  St.  Lawrence- 
Champlain  basin  ( encompassing  Quebec  as 
far  north  as  Quebec  City,  and  Vermont). 
A.  calva  is  distributed  throughout  the  Great 
Lakes  region,  but  is  not  found  in  the  Lake 
Superior  drainage  basin,  except  in  its  outlet, 
the  St.  Mary's  River.  Southward,  it  has 
been  recorded  from  the  Hudson  River  to 
western  Connecticut  ( recorded  as  the  result 
of  introduction;  Hubbs  and  Lagler,  1967); 
Harrisburg,  Pennsylvania,  to  the  Susque- 
hanna River;  and  along  the  Atlantic  slope 
to  the  Carolinas  and  Florida.  Westward, 
A.  calva  occurs  along  the  Gulf  Coast  to 
southern  Texas  as  far  south  as  Brownsville, 
and  northward,  through  eastern  Texas, 
southeastern  Oklahoma,  northwestern  Ar- 
kansas, eastern  Missouri,  and  approximately 
50  miles  west  of  the  Mississippi  River  to 
Brainard,  Minnesota. 

Pleistocene  Occurrences 

Amia  calva  has  been  reported  from  only 
three  Pleistocene  localities:  (1)  Chicago, 
Illinois,  (2)  Vero  Beach,  Florida,  and  (3) 
Itchtucknee  River  deposits,  Columbia 
County,  Florida  (MCZ  9524,  9529,  9542). 
Hay  (1911:  552)  reported  "Professor  Frank 
Baker  (Chicago  Academy  of  Science)  has 
shown  me  a  considerable  part  of  the  skele- 
ton and  scales  of  a  bowfin  which  he  found 
in  the  Pleistocene  clay  near  Chicago."  A 
thorough  search  of  the  Chicago  Academy 
of  Science  collecrions  failed  to  produce  this 
specimen.  Hay  (1917,  1923)  listed  Amia 
calva  among  the  fossil  vertebrate  remains 
found  in  the  Pleistocene  sands  at  Vero 
Beach,  Florida.  Swift  ( 1968 ) ,  in  his  review 
of  fossil  fishes  of  Florida,  figured  Hay's 
Aiiiia  specimens  (left  dentary  and  a  gular 
plate;  FSM  collections)  and  concluded  that 


Fossil  Amiids  •  Borcskc 


/  /«°«'m;;; L.._  ;  , 

/■  \  r — ■'• ..—■•J^ 

!         \  :  NORTH  DAKOTA  ;  ''w 

,      L  .  I        \  I  \minnesota 

'""^GO- ^^  >  •  to  '  ■» 

^^^^^^■>.-,.    /  /o*  ^1  ! 

I  liJUH—^  m    o  fe'"   oo     ^— i 

/  /         'l-^-.-    i  '^ 

V  /  °       /'^OlORAdS"— ^  — 1  •(__ 

\  /  /■  j. VMlSbl)    I 

\  /'  ;  ;  KANSAS       '       '  "^ 

\  /  J  !  \ 

\        L  .'  ^      I        o  , 

\     .j^^'^oZ. f. !  I 

\  I  ;  "^w  MEx-6 T-«- -I 

V;  •     Q  *• .OKLAHOMA  L 

T  /  -TEXAS      ^  7; 

>  /•  I  ^ 

..    ^  /  i         '. 

Fossil    forms  of   "••v^  '  !  . 

©    /i3/77/(7  Sp.  'N 

•  A.fragosa  \  s'"~-. 

°  A.uintaensis  ''■^      '\ 

a  A.c\.uintaensis  \ 

'  A. scuta  to  *• 

»  /J.cf.  scuta  fa  \ 

■  Amiidae    incertae  sedis 

X  Pleistocene  location  of  A.calva 


Fig.    1.      Distribution    of   km\a    calva.    Fossil    occurrences   of   Amia    spp.    ore    represented    by    symbols    explained    in    the 
legend. 


A.  calva  was  probably  very  common  in  the 
Pleistocene  fresh  waters  of  the  United 
States.  The  pancity  of  Pleistocene  material 
does  not  necessarily  mean  the  fish  was  not 
common  in  the  Pleistocene,  but  does  indi- 
cate that  A77ua  remains  have  not  been 
extensively  collected  or  identified  in  exist- 
iii'j;  museum  Pleistocene  collections. 

Diagnosis 

Vertebral  meristics  similar  to  A.  sctitata, 
but  head/standard-length  proportion  (0.271 
mean)  is  smaller  than  in  the  fossil  forms. 
Extrascapular  strap-shaped  and  relatively 
wide  at  midlitie,  as  in  A.  scutata,  but  pos- 
terior edge  is  curved  so  that  it  is  proximally 
convex,  then  concave  toward  the  distal 
corner,  which  results  in  a  posterolateral 
projection.  Pterotic  borders  frontal  pos- 
teriorly rather  than  laterally;  anterior  end 
is  as  wide  as  posterior  end.  Orbital  excava- 
tion is  shallower  than  in  other  species,  with 


a  mean  depth-to-length  ratio  of  0.100.  In- 
fraorbital 4  is  smaller  than  infraorbital  5, 
less  robust  than  in  fossil  Amia.  Preopercu- 
lum  as  wide  dorsally  as  ventrally.  Symphy- 
seal  incurx'ing  of  dentary  relatively  less  than 
ill  A.  fraii^osa,  ])ut  greater  than  in  A.  scutata 
and  A.  uintacnsis:  little  or  no  overlapping 
of  dorsal  coronoid  articulation  surface  on 
ventral  siuface  of  ramus;  deep  Meckelian 
groov^e.  Ventropostcrior  process  of  cleith- 
rum  less  sculptured  than  in  other  species  of 
Amia.  V^omerine  teeth  shaip,  conical,  num- 
bering between  15-27,  more  anteriorly 
placed  than  in  A.  uintacnsis  or  A.  fraii.osa. 
Bones  less  ossified  than  in  fossil  Amia. 
Greatest  known  standard-length  650  mm. 

MORPHOMETRICS 

Comparison  of  morphometric  and  meris- 
tic  data  of  Recent  and  fossil  Amia  has 
facilitated  an  e\'aluation  of  the  taxonomy  as 
well  as  clarified  anatomical  trends.    Many 


6        Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


generic  or  specific  character-states  for  Amia 
''dicfyocepJmla,'^  Amia  scutata,  "Paramiatus 
gurleiji,^'  and  Amia  fragosa  have  been  pre- 
viously estabhslied  on  osteological  data 
based  primarily  on  gross  anatomical  propor- 
tions (head/standard-lcngth  ratio  and  posi- 
tions of  insertion  of  pelvic  and  anal  fins/ 
standard-length  ratios)  and  skull  propor- 
tions (parietal/frontal  and  operculum 
width/length  ratios).  Meristic  character- 
states  have  also  been  used  for  A.  "dictijo- 
cephala"  and  A.  scutata. 

Altliough  an  age-growth  analysis  on  Amia 
calva  was  done  by  Cartier  and  Magnin 
(1967),  no  moiphometric  investigation  of  a 
growth-series  of  Recent  A.  colva  has  yet 
been  completed  or  used  for  comparison 
with  fossil  forms.  Estes  and  Berberian 
(1969:  10)  suggest  that  knowledge  of  the 
growth-series  of  A.  calva  would  be  of  con- 
siderable importance  in  tracing  the  ancestry 
of  the  modern  species. 

Hammett  and  Hammett  (1939)  made  a 
moi-phometric  study  of  the  Recent  Lepisos- 
teus  platijrhinciis,  taking  length  dimensions 
of  a  sample  of  live  fish  from  Florida.  Since 
Lepisosteus,  like  Amia,  is  one  of  the  two 
extant  holosteans,  their  analysis  is  poten- 
tially useful  in  providing  information  on  the 
ancient  species.  However,  they  did  not 
actually  compare  the  live  material  or  data 
with  any  fossil  material. 

According  to  Imbrie  (1956),  Simpson 
et  al.  (1960),  and  Gould  (1966),  growth 
studies  offer  excellent  means  with  which  to 
clarify  evolutionary  and  taxonomic  prob- 
lems in  the  fossil  record.  An  interesting 
model  utilizing  morphometric  data  for 
synonymy  of  fossil  forms  was  made  by 
Thomson  and  Hahn  ( 1968 )  on  the  growth- 
series  patterns  of  Devonian  rhipidistian 
fishes,  in  which  they  showed  that  Thiirsius 
clappi  was  actually  a  juvenile  form  of 
Eusthenopteron  foordi.  In  studying  fossil 
material,  as  Thomson  and  Hahn  (196S: 
201)  indicate,  there  is  a  problem  in  deter- 
mining the  age,  sexual  matmity,  and  envi- 
ronmental regime  of  the  animal.  Also,  of 
course,  it  is  necessary  to  have  sufficient  fos- 


sil material  with  which  to  erect  an  adequate 
growth-series. 

This  present  analysis  is  undertaken  (1) 
to  determine  w  hether  skull  and  axial  skele- 
tal proportions  of  amiids  are  isometric  or 
allometric  with  increasing  size,  (2)  to 
establish  the  variation  in  meristic  charac- 
ters of  Recent  A.  calva,  and  ( 3 )  to  compare 
moiphometrics  and  meristics  of  Recent  A. 
calva  with  those  of  the  fossil  forms.  This 
study  utilizes  a  small  sample  of  18  Recent 
A.  calva  specimens  from  the  St.  Croix  River, 
Wisconsin.  Measurements  were  taken  from 
a  growth  series  that  includes  the  size  range 
of  most  of  the  articulated  fossil  forms.  The 
largest  A.  calva  specimen,  from  St.  Joseph 
County,  Michigan  (UMMZ  197683),  was 
analyzed  to  see  whether  the  large  specimen 
would  agree  with  the  anatomical  propor- 
tions and  meristic  characters  of  the  Wiscon- 
sin specimens.  Three  smaller  specimens 
from  Pewaukee,  Wisconsin  (MCZ  8970), 
were  also  included.  The  fossil  sample  con- 
tains six  complete  and  ten  partially  com- 
plete amiid  specimens  ranging  in  age  from 
Late  Cretaceous  to  Late  Miocene  which, 
although  moiphometrically  similar  in  vary- 
ing degree,  are  too  few  to  warrant  conclu- 
sions in  themselves. 

Methods 

Measurements  chosen  for  this  study  ( Fig. 
2)  are  those  of  Hubbs  and  Lagler  (1967: 
20).  In  fossil  forms,  because  of  the  lack  of 
preservation  of  internal  soft  anatomy  as  well 
as  the  impossibility  of  determining  their 
interbreeding  potential,  these  particular 
measurements  necessarily  assume  an  in- 
creased taxonomic  significance,  since  they 
often  provide  the  only  viable  parameters 
with  which  to  designate  genera  and  species. 
Measurements  taken  on  A.  calva  are  limited 
to  those  also  represented  in  the  fossil  speci- 
mens. Each  of  the  A.  calva  measured  was 
X-rayed,  except  for  three  small  specimens, 
which  were  cleared  and  stained.  The  range 
of  error  for  aj]  measmements  taken  on  Re- 
cent and  fossil  material  is  ±0.04  mm.  The 
range  of  error  for  the  ratios  is  ±0.08  mm; 


Fossil  Aaiiids  •  Boreske 


Fig.   2.      Index  to  the   measurements   used,   superimposed   upon   an   outline   drawing  of  Amia. 


Key  for  body  measurements: 

TL    =  Total-Length 

SL    =z  Standard-Length 

H     zi:  Head-Length 

C     ^  Caudal-Length 

Pf    =3  Insertion    of    Pelvic    Fin 

P     =  Insertion    of   Anal    Fin 

HL  =:  Standard-Length   minus   Head-Length 

ML  :=  Standard-Length   minus  Mandible-Length 


Key  for  abbreviations  of  cranial  elements  used  in  morphometric  study: 


M 

= 

Mandible 

G 

Gular 

|5 

Infraorbital 

F 

= 

Frontal 

Par 

= 

Parietal 

O 

zi: 

Operculum 

Table  1.     Length  dimensions  of  22  specimens  of  Amia  calva  L.:  21  from  Wisconsin 

(MCZ  8970'),  1  from  Michigan  (  UMMZ  197683)'* 

Measurements  in  mm 


Specimen 

Class  Range 

Code 

Total  Length 

No. 

TL 

SL 

ML 

H 

HL 

Pf 

P 

c 

1* 

80.0 

1 

80.0 

70.5 

57.0 

22.0 

48.5 

32.5 

12.5 

10.5 

2* 

95.0-105.0 

2 

100.0 

85.0 

70.0 

25.0 

60.0 

39.5 

16.0 

15.0 

3 

207.0-212.0 

6 

210.0 

175.0 

145.0 

50.5 

124.5 

80.9 

35.0 

35.0 

4 

227.9-232.0 

4 

230.0 

193.0 

161.0 

54.6 

138.4 

88.5 

35.5 

36.0 

5 

241.0 

241.0 

199.0 

165.9 

56.8 

142.2 

93.5 

38.0 

42.0 

6 

291.0 

291.0 

237.0 

197.0 

64.0 

173.0 

115.0 

52.5 

54.0 

7 

310.0 

310.0 

248.0 

207.0 

68.5 

179.5 

112.0 

46.0 

62.0 

8 

339.0 

339.0 

274.0 

230.0 

73.0 

202.0 

125.0 

51.0 

64.0 

9 

385.0 

385.0 

313.0 

259.0 

82.0 

231.0 

142.0 

52.0 

72.0 

10 

433.0 

4.33.0 

349.0 

293.0 

91.0 

258.0 

170.0 

71.0 

84.0 

11 

475.0 

475.0 

.399.0 

335.0 

103.0 

296.0 

181.0 

82.0 

76.0 

12 

507.0 

507.0 

423.0 

359.5 

109.0 

317.0 

192.0 

93.0 

81.0 

13»» 

756.0 

756.0 

648.0 

545.3 

164.0 

480.0 

299.0 

138.0 

102.0 

See  Figure  2  for  abbreviations. 


8         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


this  margin  of  error  is  graphically  inconse- 
quential in  this  study.  Specimens  of  A.  calva 
whose  total  length  was  between  207  mm 
and  507  mm  were  selected  because  this 
range  of  A.  calva  would  provide  the  best 
information  for  comparison  with  the  fossil 
species.  Twenty-two  specimens  of  A.  calva 
were  measured  (Table  1).  Eighteen  of 
these  are  from  the  St.  Croix  River,  Wiscon- 
sin. These  18  specimens  of  A.  calva  fall 
into  ten  categories  arranged  here  by  ap- 
proximately 20-30-mm  class  range  incre- 
ments in  total-length.  Although  these 
categories  represent  arbitrary  rather  than 
biological  growth  stages,  they  provide 
suflBcient  information  on  the  morphologic 
size  changes  of  A.  calva.  Three  smaller 
specimens  (MCZ  8970,  also  from  Wiscon- 
sin) witli  a  size  range  of  80-105  mm 
total  length  (TL)  were  included  to  de- 
termine whether  they  would  follow  the 
predicted  allometric  effect  on  the  growth- 
series  Hne,  since,  as  Thomson  and  Hahn 
(1968:  205)  note,  it  is  a  common  feature 
for  the  early  stages  of  juvenile  animals  to 
have  heads  proportionately  larger  than  the 
bodies.  Hay  (1895)  notes  that  an  80-mm 
A.  calva  is  beyond  the  embryonic  stage  and 
is  an  early  juvenile  with  most  of  its  bones 
at  least  partially  ossified.  The  80-mm  speci- 
men has  a  proportionately  larger  head  to 
standard-length  ratio  than  the  other  mem- 
bers of  the  growth-series  (Table  3).  Al- 
though this  ratio  decreases  slightly  with 
increasing  size,  the  head /standard-length 
ratio  of  0.312  for  the  80-mm  specimen  does 
not  deviate  far  from  the  growth-series  line 
(Figs.  3-4). 


The  largest  specimen  (UMMZ  197683) 
was  used  as  a  size  limit  for  the  other  end 
of  the  growth-series  continuum.  It  may  be 
assumed  that  this  fish  had  already  reached 
the  size  or  point  of  maturity  at  which  fish 
normally  begin  to  decrease  their  rate  of 
growth.  This  specimen  still  retains  the 
morphological  proportions  of  the  smaller 
specimens  (Figs.  3-4)  and,  like  them,  falls 
remarkably  close  to  the  constant  relative 
size-growth  lines  of  the  various  proportions. 
Although  from  Michigan,  this  specimen 
does  not  appear  to  deviate  from  the  growth- 
series  line  established  by  the  Wisconsin 
specimens  of  A.  calva.  The  Michigan  speci- 
men of  A.  calva,  since  it  agrees  with  the 
growth-series  continuum  established  by  the 
Wisconsin  specimens,  is  helpful  in  extend- 
ing comparison  to  the  larger  fossil  amiids: 
"Paratniatus  gurleiji"  (FMNH  2201),  Amia 
fra^osa  (MCZ  5341),  and  Amia  uintaensis 
(PU  13865),  which  are  outside  the  size 
range  of  the  Wisconsin  sample. 

General  Proportions  and  Growth 

Allometric  growth,  according  to  Gould 
(1966:  595),  describes  geometrically  pro- 
gressive change  in  shape  or  proportions 
with  size,  and  is  generally  represented  by  a 
curvilinear  line  or,  in  certain  cases,  by  a 
straight  line  in  which  the  Y-intercept  is 
significantly  different  from  0. 

For  the  Amia  calva  growth  series  dis- 
cussed here,  the  ordered  pairs  correspond- 
ing to  the  proportions  in  each  series  have 
been  plotted  on  a  graph,  as  well  as  the 
straight  line  corresponding  to  the  equation 
y  =  a  +  bx  (of  the  best  fit  computed  ac- 


Table  2. 

Length 

DrMENSIONS 

;   OF   6   ARTICULATED  FOSSIL 

AMIIDS 

Measurements  in 

mm 

TL 

SL 

ML 

H 

HL 

Pf 

P 

C 

A.  scutata  PU  10172 

'■«yo4.o 

339.0 

276.5 

106.0 

233.0 

159.0 

73.0 

e«t65.0 

A.  scutata  UMMP  V-57431 

— 

388.0 

313.8 

121.0 

267.0 

183.0 

83.0 

A.  kehreri  BMNH  P33480 

249.0 

191.0 

160.8 

59.2 

131.8 

89.0 

38.5 

58.0 

"Paramiatus  gurleiji" 

FMNH  2201 

702.0 

510.0 

430.0 

157.0 

353.0 

<'«f265.0 

78.0 

192.0 

A.  fragosa  MCZ  5341 

575.0 

455.0 

383.0 

142.0 

313.0 

210.0 

75.0 

115.0 

A.  uintaensis  FV   13865 

848.0 

664.0 

— 

214.0 

450.0 

288.0 

116.0 

160.0 

See  Figure  2  for  abbreviations. 


Fossil  Amhds  •  Boreske        9 


320- 


280- 


240- 


200- 


160- 


120- 


80- 


40- 


I 

40 


—I 1         I         I         I 

120  200  280 

STANDARD 


I  I  I 

360 
LENGTH     mm 


680 


440 


520 


600 


Fig.  3.  Relative  growth-lines  of  head-length  (H),  pelvic  fin  insertion  (Pf),  and  ana!  fin  insertion  (P)  plotted  arith- 
metically against  standard-length,  for  18  specimens  of  Recent  Ami'o  calva  (A  =  MCZ  8970  and  H  =  UMMZ 
197683   are   included   for  comparison). 

cording  to  the   method  of  least  squares);  whicli  nearly  passes  through  the  origin  of 

the  results  of  these  calculations  appear  in  the  graph.    The  coefficient  of  correlation  is 

Figures  3-4.    Practically   all   the   ratios   in  almost  equal  to  1.0  in  each  case,  an  indica- 

Figure  3  fall  onto  straight   lines,   each   of  tion  that  the  computed  straight  line  provides 


10        Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Table  3.     Comparisox  of  length  proportions  in  22  specimens  of  Amio  calva  with  fossil  amiids 


Specimen 

Code 

H/SL 

Pf/SL 

P/SL 

1 

0.312 

0.461 

0.177 

2 

0.294 

0.464 

0.188 

3 

0.289 

0.462 

0.200 

4 

0.283 

0.459 

0.184 

5 

0.285 

0.470 

0.191 

6 

0.270 

0.485 

0.222 

7 

0.276 

0.452 

0.185 

8 

0.266 

0.456 

0.186 

9 

0.262 

0.455 

0.198 

10 

0.261 

0.487 

0.203 

11 

0.258 

0.454 

0.206 

12 

0.258 

0.454 

0.220 

13 

0.259 

0.461 

0.213 

0.258-0.289'' 

0.452-0.487* 

0.184-0.222* 

mean  =  (0.271)* 

mean  =  (0.463)' 

mean  =  (0.199)* 

Oligocene 

A.  scutata  PU 

10172 

0.313 

0.469 

0.215 

A.  scutata  UMMP  V-57431 

0.312 

0.472 

0.214 

Eocene 

A.  kehreri  BMNH  P33480 
"Paramiatus  gurleyi" 

FMNH  2201 
A.  fragosa  MCZ  5341 
A.  uintaensis  PU  13865 


0.310 


0.466 


0.201 


0.308 

*'sto.520 

0.153 

0.312 

0.462 

0.165 

0.322 

0.434 

0.175 

Range  and  mean  exclude  Specimen  Codes  1  &  2  (MCZ  8970)  and  13  (UMMZ  197683; 


a  very  good  fit  for  the  ratio  series,  and  that 
the  relative  growth  of  these  three  dimen- 
sions is  essentially  isometric  rather  than 
allometric.  The  Wisconsin  specimens  (in- 
cluding the  80-105-mm  specimens )  and  the 
larger  Michigan  specimen  all  fall  close  to 
the  line  calculated  for  each  of  the  three 
ratios  (Fig.  3).  The  proportions  of  head- 
length/standard-length,  insertion  of  pelvic 
fins/standard-length,  and  insertion  of  anal 
fins /standard-length  are  shown  in  Table  3. 
The  head/standard-length  ratio  shows  a 
slight  decrease  with  increasing  size,  but 
this  ratio  series  nonetheless  has  a  very  high 
coefficient  of  correlation  for  the  strength  of 
the  linear  relationship  (Fig.  4). 

The  lengths  of  the  mandible,  parietal, 
frontal,  and  operculum  in  Recent  A.  calva 
appear  in  Table  4,  and  the  proportional 
ratios  in  Table  6.  The  relative  growth  rate 
of  each  of  these  proportions  is  constant  with 
X  and  Y-intercepts  of  the  straight  line  close 
to  the  origin.  The  coefficient  of  correlation 


for  the  variables  in  each  of  the  proportions 
is  0.997,  0.975,  and  0.997,  respectively  ( Fig. 
5).  Combined,  these  two  factors  indicate 
constant  and  therefore  isometric  relative 
size-growth  of  the  compared  skull  element. 

Comparisons  with  Fossil  Forms 

Six  articulated  fossil  specimens  were 
available  for  measurement  and  calculation 
of  head /standard-length  and  positions  of 
insertion  of  pelvic  and  anal  fins/ standard- 
length  (Tables  2-3).  The  measurements 
taken  from  the  fossil  forms  are  as  exact  as 
conditions  allow,  although  it  must  be 
stressed  that  varying  degrees  of  crushing 
and  distortion  have  occurred  in  fossilization, 
and  evaluation  of  the  morphometries  should 
be  qualified  with  this  in  mind. 

Head /standard-length  ratios  (Fig.  4). 
The  fossil  forms  all  show  a  slightly  greater 
head/ standard-length  ratio  than  does  the 
Recent  species  (Table  3;  Fig.  4).  A. 
uintaensis    (PU    13865)    is   the   largest   of 


Fossil  Amiids  •  Boreske 


11 


Table  4.     Length  dimensions  of  Mandible  (M),  Gular  (C),  Frontal  (F),  Parietal  (Par), 
Infraorbital  ^(I''),  and  Operculum  (O)  in  22  specimens  of  A.  calva 

Measurements  in  mm 


Operci 

ilum 

Specimen 

Dors. -Vent. 

Ant.-Post. 

Code 

M 

G 

F 

Par 

18 

(OL) 

(OD) 

1* 

13.5 

8.0 

10.0 

5.0 

5.0 

8.1 

7.5 

2* 

14.9 

9.4 

11.4 

6.1 

5.8 

9.0 

8.4 

3 

30.0 

19.0 

18.0 

9.7 

12.0 

14.0 

12.9 

4 

32.0 

21.0 

20.0 

9.9 

13.5 

15.0 

14.2 

5 

33.5 

20.5 

19.0 

10.0 

14.2 

16.0 

14.9 

6 

39.0 

26.0 

25.0 

11.0 

17.0 

16.9 

16.5 

7 

42.0 

28.0 

25.5 

11.5 

18.0 

18.1 

17.5 

8 

45.0 

27.0 

26.2 

13.5 

21.0 

19.8 

18.5 

9 

53.0 

31.0 

31.0 

14.7 

22.5 

22.8 

21.8 

10 

56.0 

32.5 

32.2 

17.0 

26.5 

22.6 

22.5 

11 

63.0 

39.0 

38.4 

18.5 

30.5 

28.1 

27.8 

12 

66.5 

42.5 

39.0 

19.5 

31.5 

27.8 

28.5 

13** 

102.7 

— 

60.7 

30.0 

— 

— 

o  MCZ  8970. 
«<»  UMMZ   197683. 


all  the  fossil  specimens,  but  nonetheless 
has  a  greater  head/ standard-length  ratio 
than  any  of  the  others.  The  head  of  this 
form  is  so  much  more  elongated  than  the 
head  in  A.  fra^oso  (MCZ  5341),  A.  kehreri 


(BMNH  P33480),  and  "Paramiatus  gurleiji" 
(FMNH  2201)  that  it  offsets  the  fact  that 
its  vertebral  column  includes  approximately 
20  more  vertebrae  than  do  these  three  forms 
(Table  9).    Thus,   although  A.   idntaensis 


Table  5.     Length  dimensions  of  Head  (H),   Mandiiu^  (M),  Gular  (G),  Frontal  (F), 
Parietal  (Par),  Infraorbital  ^'{V'),  and  Operculum  (O)  in  fossil  amiids 

Measurements  in  mm 


Operculum 

Dors.-Vent. 

Ant.-Post. 

H 

M 

G 

F 

Par 

F 

(OL) 

(OD) 

A.  cf.  scutata  UCMP  38222 

— 

65.2 

— 

46.0 

23.0 

35.0 

— 

— 

A.  scutata  PU  10172 

106.0 

62.5 

31.2 

35.0 

16.0 

— 

29.0 

28.0 

A.  scutata  UMMP  V-57431 

121.0 

74.2 

44.3 

20.0 

29.1 

27.9** 

A.  "dictt/ocephala" 

AMNfl  2802 

111.5 

68.0 

38.0 

17.0 

29.1 

32.0 

30.0 

A.  kchrcri  BMNH  P33480 

59.2 

30.2 

— 

20.0 

8.4 

15.3 

20.5 

19.0 

"Paramiatus  fiurletji" 

FMNH  2201 

157.0 

80.0 

— 

58.0 

23.6 

25.0 

40.0 

37.0 

A.  fra^osa  MCZ  5341 

142.0 

72.0 

68.7 

56.0 

22.8 

39.0 

36.2 

A.  fragosa  MCZ  9264 

80.0 

40.0 

— 

26.0 

10.5 

18.5 

— 

— 

A.  uintaensis  PU  13865 

214.0 

— 

— . 

88.0 

34.0 

— 

55.0 

51.0 

"Protamia"  mongoliensis 

AMNH  6372 

— 

— 

81.0 

— 

— 

— 

54.0 

52.0 

A.  uintaensis  PU  16236 

315.0** 

220.0 

158.0 

160.0 

60.0 

— 

— 

95.0 

A.  fragosa  MCZ  9291 

— ■ 

— 

— 

— 

— 

— 

27.0 

25.0 

A.  fragosa  AMNH  9315 

— 

— 

— 

— 

29.0** 

27.0 

A.  fragosa  UA  5450* 

— 

— 

26.0 

10.0 

— 

— 

— 

A.  fragosa  UA  5458* 

— 

— 

30.0 

12.0 

— 

— 

A.  fragosa  UA  5480* 

— 

— 

— 

20.0 

26.0 

24.0 

•Data   from   O'Brien    (1969). 
••  Est. 


12        Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.   1 


240- 

220- 
200- 

EQUATiONof  STRAIGHT  LINE-  M=  7.268  +  (0,234)  (SLl 

u 
y 

leo- 

COEFFICIENT  ol  CORRELATION  =  0.999 

ISO- 

MEAN   ^  -.  0.271 

STANDARD  DEVIATION  5^  =  0.0117 

COEFFICIENT  of  VARIATION  =4.33% 

E 

E   140. 

X 

;  120- 

a 

S  100- 

I 

^^ 

60- 

xX' 

SO' 

/^ 

40' 

/^ 

20- 

^x 

0. 

320  400  460 

STANDARD     LENGTH    mm 


y 

EQUATION  of  STRAIGHT  LINE  -  PF  =  1  438  +  (0.454)(SL) 

■^ 

288  ' 
240- 

COEFFICIENT  of  CORRELATION  .0.995 
MEAN   f^  '-  0.463 
STANDARD  DEVIATION  |^  -■  00131 
COEFFICIENT  of  VARIATION  ■  2  83% 

y  '" 

0                 y 

y 
y 
y 
y 

y 

192- 

y 

144. 

96- 

y 

48. 

X 

0. 

X 

240  J20  400 

STANDARD     LENGTH    mm 


E 
E 
■z-     120' 


EQUATION  of  STRAIGHT  LINE  ■■  P  =     -5.171   +(O.E27)(SL) 

COEFFICIENT  of  CORRELATION  =  0.987 

MEAN  ^  =  0.199 

STANDARD  DEVIATION^  .0.0135 

COEFFICIENT  ot   VARIATION  '6.78  % 


240       320       400 
STANDARD  LENGTH  mm 


Fossil  Ami  ids  •  Boreske        13 


has  significantly  more  vertebrae  than  these 
other  forms,  this  feature  is  not  reflected  in 
a  comparison  of  head/ standard-length  ra- 
tios ( Table  3 ) .  This  is  also  true,  to  a  lesser 
extent,  in  both  A.  scutata  specimens   (PU 
10172,  UMMP  V-57431)  from  the  Ohgocene 
Florissant  Formation;  these  specimens  fall 
into  the  head/ standard-length  range  of  the 
three    fore-mentioned    forms,    but    like    A. 
uintaensis,  possess  vertebral  columns  having 
nearly  the  same  number  of  centra  as  those 
in  A.  calva.   Thus  in  themselves  the  head/ 
standard-length  ratios  are  of  little  help  in 
comparing    the    fossil    forms,    but    when 
coupled  with  the  corresponding  lengths  of 
the   vertebral   column    (based   on   number 
of  centra)  they  are  informative.  A.  uintaen- 
sis (PU  13865)  and  A.  scutata  (PU  10172, 
UMMP  V-57431)  have  relatively  elongated 
heads;    A.    kehreri    (BMNH    P33480),    A. 
fragosa  (MCZ  5341),  and  "Paramiatiis  g,ur- 
leyi"  (FMNH  2201)  have  relatively  shorter 
heads,  since  the  head/standard-length  ratio 
is  less  than  might  otherwise  be  expected 
considering    the    smaller    total-number    of 
centra    (only    two-thirds    the    number    of 
centra  of  A.  uintaensis,  A.  scutata,  and  A. 
calva ) .  A  tentative  growth-line  ( also  calcu- 
lated by  the  best-fit  method )  was  included 
for  A.  jragosa  on  the  basis  of  three  speci- 
mens   (Fig.   4).    In   comparison   with   the 
growth-line   of  the   Recent   species    (0.271 
mean),  it  reflects  the  larger  head/ standard- 
length  ratio  of  A.  fragosa    (0.310  mean). 
The  growth-line  computed  for  A.  jragosa 
is  linear  and  falls  near  the  origin,  indicating 
that  increase  in  head  size/ standard -length 
was  isometric,  as  in  A.  calva. 

Fin  relations] lips.  In  the  smaller  fossil 
forms,  the  ratio  of  the  point  of  insertion 
of  the  pelvic  fin/ standard-length  shows  little 
deviation  from  the  modern  species  (Table 
3;  Fig.  4)  except  for  two  Eocene  specimens, 
"Paramiatus  gurleyi"    (FMNH   2201)    and 


A.  uintaensis  (PU  13865),  which  fall  out- 
side of  the  range  on  either  side  of  tlie  size- 
growth  line.  The  greater  ratio  for  "Para- 
miatus gurleyi,"  however,  is  probably  the 
result  of  distortion  in  its  preservation.  The 
length  of  the  pelvic  fin  insertion/ standard- 
length  does  not  appear  to  be  a  satisfactory 
taxonomic  index,  distinguishing  neither  the 
fossil  forms  from  one  another  nor  the  fossil 
forms  from  the  Recent  A.  calva. 

"Paramiatus  gurleyi"  (FMNH  2201),  A. 
uintaensis  (PU  13865),  and  A.  jragosa 
(MCZ  5341)  have  a  relatively  shorter 
dimension  between  the  anal  fin  and  the  end 
of  the  vertebral  column  than  do  A.  calva, 
A.  scutata,  and  A.  kehreri  (Fig.  31).  Any 
attempt  to  inteipret  the  fossil  data  for  this 
ratio  is  complicated  by  the  fact  that  con- 
siderable overlap  with  the  Recent  species 
occurs.  Both  long-bodied  (A.  scutata)  and 
short-bodied  (A.  kehreri)  forms  fall  within 
the  range  of  A.  calva,  while  other  long- 
bodied  (A.  uintaensis)  and  short-bodied 
(A.  jragosa,  including  "Paramiatus  gur- 
leyi") fonns  fall  below  the  range  of  the 
Recent  species  (Table  3).  Although  the 
ratio  of  anal  fin/ standard-length  may  pos- 
sibly be  useful  in  distinguishing  A.  jragosa 
(including  "Paramiatus  gurleyi")  from  A. 
calva,  A.  scutata,  and  A.  kehreri,  it  is 
not  useful  in  distinguishing  either  of  the 
two  fossil  fonns  from  one  another  or  from 
A.  calva.  The  smaller  dimension  indicated 
by  the  low  ratios  (0.153,  0.165)  of  A. 
jragosa  is  doubtless  a  reflection  of  its 
shorter  axial  column.  The  relatively  small 
(0.175)  ratio  for  A.  uintaensis  is  probably 
in  part  the  result  of  its  longer  head,  wliich 
increases  its  standard-length  in  relation  to 
the  other  forms;  at  any  rate,  the  difference 
between  the  A.  uintaensis  ratio  and  the 
range  for  Recent  A.  calva  is  not  very  sig- 
nificant. 

Mandible  I  head  ratios.     A  comparison  of 


Fig.  4.  Relative  growth-lines  (broken-solid  lines)  of  head-length,  pelvic  fin  insertion,  and  anal  fin  insertion  plotted 
arithmetically  against  standard-length  for  Recent  Amia  calva  (A  =  MCZ  8970  and  ■  =  U/AMZ  197683  are  included 
for  comparison)  with  compared  fossil  forms:  fl  =  A.  hagosa  (A.  kehreri)  BMNH  P33480;  f-  =  A.  fragosa  (Pararr^iafus 
gurleyi)  FMNH  2201;  f'-^  =  A.  fragosa  MCZ  5341;  s^  =  A.  scufafa  PU  10172;  s^  =  A.  scutofo  UMMP  V-57431; 
u  =  A.    uintaensis   PU    13865.   The   broken-dotted   line   is  the   "best  fit"    line   for  available   specimens   of   A.   fragosa. 


14         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


no 

100 
90^ 


tOuATiON  o(  STRAIGHT   LINE  ■  M  ■-!  010  +  (0-636HM) 

COEFFICIENT  of   CORRELATION  ^  0.  997 

MEAN  ^  '  0  609 

STANDARD   DEVIATION   ^  ■  0  0171 

COEFFICIENT  of  VARIATION  -■  2.81  % 


55- 

50. 

/ 

45- 

EQUATION  0)  STRAIGHT  LINE  ^  PAR  =  0  278 +  (0  479HF)             '^                    y 
COEFFICIENT  ol  CORRELATION  •  0.975                                       /^                    ^ 

40- 

E 

MEAN  ^  .  0.495                                                                        /                  /"^ 

E 
± 

35- 

STANDARD   deviation""-  0  0333                                ^^                 / 
COEFFICIENT  of  VARIATION- 6  73%                           /                 /  "^  V 

O 

/                / 

X 
UJ 

30- 

f'  y 

~t 

/  / 

< 

/  / 

UJ 

25- 

/ 

< 

V  A 

S 

/    /•' 

20- 

1!- 

10. 

/^' 

5. 
0. 

/ 

30         40  50         60 


SO        90         rOO       110       120 


FRONTAL     LENGTH    mm 


o-»r 


EQUATION  of  STRAIGHT  LINE  :  OD  ■  - 1.559  +  (l.079)(&LI 

COEFFICIENT  of  CORRELATION  -  0.997 

MEAN  g2  -0.964 

STANDARD  DEVIATION   ^  -  0.0326 

COEFFICIENT  of  VARIATION-  3.40% 


20  30  40 

OPERCULUIM     LENGTH  mni 


Fossil  Amiids  •  Boreske 


15 


Table  6.     Cranial  proportions  in  22  specimens 
OF  A.  calva 


Specimen 

Code 

M/H 

Par/F 

OD/OL 

1 

0.614 

0.500 

0.926 

2 

0.596 

0.535 

0.933 

3 

0.594 

0.539 

0.921 

4 

0.586 

0.495 

0.947 

5 

0.590 

0.526 

0.931 

6 

0.609 

0.440 

0.976 

7 

0.613 

0.451 

0.967 

8 

0.616 

0.515 

0.934 

9 

0.646 

0.474 

0.956 

10 

0.615 

0.528 

0.996 

11 

0.611 

0.482 

0.989 

12 

0.610 

0.500 

1.025 

13 

0.626 

0.497 

0.586- 

0.440- 

0.921- 

0.646* 

0.539" 

1.025" 

mean 

mean 

mean 

=  (0.609)" 

=  (0.495)" 

=  (0.964)" 

■*  Range    and    mean    exclude    Sjiecimen    Codes     1     &     2 
(MCZ  8970)   and   13    (UMMZ    197683). 

the  mandible/head  ratios  of  Recent  A.  calva 
with  those  of  the  fossil  forms  ( Table  7;  Fig. 
5)  indicates  that  the  A.  scutata  and  A. 
''dictyocephala"  (AMNH  2802)  ratios  are 
very  close  to  those  of  A.  calva.  The  A. 
fra^osa  .specimens  (including  "Taramiatus 
^urleijr  FMNH  2201  and  A.  kehreri  BMNH 
P33480)  have  a  mean  mandible/ head  ratio 
of  0.507,  which,  when  compared  to  the 
A.  calva  mean  ratio  of  0.609,  indicates  a 
relatively  smaller  mandible  to  head  size 
(Table  7).  Unfortunately,  A.  uintaensis 
(PU  13865)  cannot  be  used  in  this  com- 
parison, since  the  mandibles  are  buried  in 
matrix.  A  reconstruction  of  a  disarticulated 
A.  uintaensis  (PU  16236)  specimen  from 
the  Late  Pal  eocene  has  been  made,  and  its 
ratio  is  approximately  0.693.  Thus  man- 
dible/head proportions   may   be   valid   for 


distinguishing  specimens  of  A.  fra^usa  and 
A.  uintaensis  from  one  another  as  well  as 
from  A.  calva  and  A.  scutata.  This  ratio, 
however,  caimot  be  used  as  a  valid  criterion 
for  distinguishing  A.  scutata  from  A.  calva. 
The  0.693  mandible/ head  ratio  of  A. 
uintaeims  indicates  that  this  form  has  the 
largest  mouth  gape  of  the  four  valid  species. 
A  tentative  growth-line  for  the  mandible/ 
head-length  proportion  of  A.  fra<i^osa,  estab- 
lished on  four  specimens,  shows  that  its  jaw 
is  16  percent  smaller  than  that  of  A.  calva, 
and  in  this  respect  confirms  Romer  and 
Fryxell's  (1928)  reconstruction  of  "Paramia- 
tus  ^urleiji." 

Parietal / frontal  ratios.  Only  articulated 
frontals  and  parietals  were  measured  for 
this  study.  The  frontal-length  was  taken 
from  the  anteriormost  extent  of  the  dermal 
sculpture  to  the  median  point  between  the 
most  anterior  and  posterior  extents  of  the 
frontal-parietal  suture;  the  parietals  were 
also  measured  by  their  midline  anteropos- 
terior length.  The  parietal/ frontal  ratio  of 
the  fossil  forms  as  compared  with  that  of 
the  Recent  A.  calva  indicates  that,  in  vary- 
ing degree,  the  fossil  species  have  relatively 
shorter  parietals  and  longer  frontals  ( Table 
7;  Fig.  5).  The  largest  specimen  of  A. 
uintaensis  (PU  16236)  is  mostly  disarticu- 
lated, but  fortunately  the  skull  table  is  in- 
tact. It  has  the  smallest  parietal/ frontal 
ratio  of  all  the  fossil  .species,  0.375.  The 
Eocene  A.  uintaensis  specimen  (PU  13865) 
has  a  slightly  larger  ratio  of  0.386,  which 
is  nearly  equal  to  the  Edmonton  A.  jragosa 
(UA  5450).  All  the  other  available  A. 
fragosa  specimens,  including  "Paramiatus 
gurleyi'  ( FMNH  2201 ),  have  slightly  larger 
ratios  and  are  quite  consistent,  ranging  only 


Fig.  5.  Relative  growth-lines  (broken-solid  lines)  of  mondible-length  plotted  arithmetically  against  head-length, 
parietal-length  plotted  arithmetically  against  frontal-length,  and  operculum-depth  (anteroposteriorly)  plotted  arith- 
metically against  operculum-length  (dorsoventrally)  for  Recent  Amia  calva  (A  =  MCZ  8970  and  ^  ^^ '-"^'^'^^  197683 
are  included  for  comparison  when  element  measurements  are  available)  with  compared  fossil  forms:  f  =^  Amia 
fragosa  (A.  kehreri)  BMNH  P33480;  f-  =  A.  fragosa  (Paramiatus  gurleyi)  FMNH  2201;  f*  =  A.  frogosa  MCZ  5341; 
i^=A.  fragosa  MCZ  9264;  f"' =:  A.  frogoso  UA  5458;  f*' =  A.  fragosa  UA  5450;  f  =  A.  fragosa  AMNH  9315; 
f*^  =  A.  fragosa  MCZ  9291;  f'^  =  A.  fragosa  UA  5480;  si  =  A.  scufaia  PU  10172;  s- =  A.  scufaia  UMMP  V-57431; 
s-' =  A.  scufafa  (A.  dictyocephala)  AMNH  2802;  s'=A.  cf.  scufafa  UC  38222;  u^  =  A.  oin/oensis  PU  13865;  u- = 
A.  ointoensis  PU  16236;  m  =  A.  mongo/zensis  AMNH  6372.  The  broken-dotted  line  is  the  "best  fit"  line  for  available 
specimens  of  Amia  fragosa. 


16         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Table  7.     Comparison  of  cranial  proportions  of  recent  and  fossil  amiids 


M/H 


Par/F 


OD/OL 


Recent 

A.  calva  ( 18 )   ( Wise. ) 

Miocene 
A.  cf.  scutata  UCMP  38222 

Oligocene 

A.  scutofa  PU  10172 
A.  scufata  UMMP  V-57431 
A.  "dictyocephala" 
AMMH  2802 

Eocene 

A.  fce/jreh  BMNH  P33480 
"Paramiatus  gurleyi" 

FMNH  2201 
A.  /ragosa  MCZ  5341 
A.  /ragosa  MCZ  9264 
A.  uintaensis  PU  13865 
"Pappichthys"  mongoliensis 

AMNH  6372 

Paleocene 

A.  uintaensis  PU  16236 

Cretaceous 
A.  /ragosa  MCZ  9291 
A.  fragosa  AMNH  9315 
A.  fragosa  UA  5450 
A.  fragosa  UA  5458 
A.  fragosa  UA  5480 


0.586-0.646 
mean  =  (0.609) 


0.590 
0.613 


0.610 


0.510 


0.693* 


0.440-0.539 
mean  =  (0.495) 

0.500 

0.457 
0.451 

0.447 
0.420 


0.375 


0.385 
0.400 


0.921-1.025 
mean  =  (0.964) 


0.965 
0.959 


0.937 


0.927 


0.509 

0.407» 

0.925 

0.507 

0.408 

0.928 

0.500 

0.404 

— 

— 

0.386 

0.927 

_^_ 

___ 

0.963 

0.926 
0.931'' 


0.923 


Est 


between  0.400-0.408.  One  of  the  specific 
character-states  that  Estes  and  Berberian 
(1969:  6)  list  for  A.  fragosa  is  a  frontal- 
length  of  approximately  2.8  times  the  length 
of  the  parietals,  which  would  give  a  ratio 
of  0.357.  This  figure  is  smaller  than  that  of 
the  known  articulated  forms,  including  the 
specimens  from  the  Edmonton  and  Will- 
wood  formations.  Although  they  may  have 
placed  too  much  emphasis  on  this  specific 
character-state,  Estes  (1964),  Janot  (1967), 
and  Estes  and  Berberian  (1969)  are  justi- 
fied in  distinguishing  A.  fragosa  from  A. 
calva  on  this  basis  since  the  ratio  of  the 
fossil  form  is  smaller  than  that  of  the  Recent 
A.  calva,  whose  parietal /frontal  proportions 
have  a  mean  ratio  of  0.495,  the  frontals 
being  approximately  twice  the  length  of  the 
parietals.  A  tentative  growth-line  for 
parietal/ frontal  proportions,  established  on 
six    specimens    of    A.    fragosa,    including 


"Paramiatus  gurleyi"  (FMNH  2201)  and 
A.  kehreri  (BMNH  P33480),  illustrates  this 
difference  between  the  fossil  form  and  the 
Recent  species  (Fig.  5).  A.  kehreri  dis- 
plays slightly  larger  parietals,  with  a  ratio 
of  0.420,  but  considering  the  geographic 
and  temporal  differences  from  the  other 
A.  fragosa  specimens,  it  is  remarkably  close 
in  this  feature  to  its  North  American  rela- 
tives. The  parietal /frontal  proportions  of 
the  specimens  of  A.  "dictyocephala"  and 
A.  scutata  fall  near  the  lower  end  of  the 
0.440-0.539  range  of  A.  calva,  and  the 
Miocene  specimen  of  Amia  ( UCMP  38222), 
with  its  ratio  of  0.500,  is  very  near  the  mean 
for  A.  calva.  There  is  thus  a  definite  trend 
from  the  Cretaceous  to  the  Miocene  (and 
Recent)  toward  an  increase  in  parietal/ 
frontal  ratio.  The  A.  uintaensis  and  A. 
fragosa  specimens  have  parietal /frontal 
ratios  smaller  than  those  of  A.  calva,  while 


Fossil  Amiids  •  Boreske         17 


the  A.  ^'dictyucepluila"  and  A.  scutata  speci- 
mens are  close  to  A.  calva  in  this  propor- 
tion. There  is,  however,  enough  intra- 
specific  variation  of  parietal /frontal  ratios 
in  the  fossil  species  to  cause  an  interspecific 
overlap  of  the  various  forms,  so  that  it  is 
impossible  to  detennine  any  definitive  limits 
between  the  consecutive  fossil  species  and 
A.  calva. 

Operculum-depth  /  operculurn-lengtJi  ra- 
tios. Although  the  fossil  forms  have  a 
slightly  narrower  operculum-depth  relative 
to  their  operculum-length,  they  all  fall 
within  the  operculum  ratio  range  of  0.921- 
1.02.5  for  A.  calva  (Fig.  5).  Table  6  indi- 
cates that  with  increasing  size  in  A.  calva 
there  may  be  a  trend  from  a  narrower  to  a 
slightly  broader  operculum.  Romer  and 
Fryxell  (1928:  521)  describe  the  operculum 
of  "Paramiatus  <!,urleiji"  as  being  greater 
dorsoventrally  than  anteroposteriorly.  Al- 
though Cretaceous  and  Eocene  specimens 
of  A.  fragosa  have  operculum  ratios  ( 0.923- 
0.963)  lower  than  the  mean  (0.964) 
for  A.  calva,  they  still  fall  within  the  range 
(0.921-1.025)  of  tlie  Recent  form.  Thus  the 
variation  of  operculum  shape  within  A.  calva 
contradicts  Hussakof's  ( 1932 )  supposition 
that  the  operculum  in  "Pappichthys"  mon- 
goliensis  (with  a  ratio  of  0.963)  is  propor- 
tionately narrower  than  that  of  A.  calva,  as 
well  as  Estes  and  Berberian's  ( 1969 )  diag- 
nosis that  A.  fragosa  has  a  relatively  shorter 
operculum-length  as  compared  with  height 
than  A.  calva.  Janot  (1967)  was  also  cau- 
tious in  assigning  taxonomic  importance  to 
the  operculum  proportions  because  of  the 
great  variability  within  the  Recent  species. 
The  operculum  width /length  ratios  in  tlie 
fossil  specimens  show  little  taxonomic  sig- 
nificance, although,  as  Estes  and  Berberian 
(1969:  7)  note,  there  does  appear  to  have 
been  a  slight  temporal  trend  toward  a 
broader  operculum. 

Discussion 

The  six  relative  growth  proportions 
(Figs.  4-5)  that  were  plotted  for  the  A. 
calva  growth-series  remained  constant  and 
therefore  isometric.   This  may  be  explained 


by  the  fact  that  these  ratios  are  derived 
from  external  rather  than  internal  dimen- 
sions, and,  as  Gould  (1966)  points  out, 
it  is  usually  the  internal  elements  that  must 
increase  at  an  allometric  rate  in  order  to 
maintain  the  external  surface  area,  whose 
dimensions  may  be  increasing  at  an  iso- 
metric rate  (see  meristic  study).  It  may 
be  assumed  that  the  relative  growth  for 
these  six  proportions  also  maintained  an 
isometric  rate  in  the  fossil  forms,  since  their 
ratios  invariably  fall  near  the  growth-lines 
for  the  Wisconsin  A.  calva  sample  (Figs. 
4-5).  However,  this  assumption  would 
have  to  be  confirmed  with  an  actual 
growth-series  of  the  fossil  forms. 

The  comparison  of  Recent  with  fossil 
forms  has  also  made  it  possible  to  determine 
the  taxonomic  value  of  the  skull/ body  and 
skull  proportions.  The  moiphometric  com- 
parison of  the  fossil  forms  with  Recent  A. 
calva  suggests  the  following  taxonomic  and 
phylogenetic  trends : 

1.  All  the  fossil  forms  have  slightly  longer 
heads  relative  to  their  standard-length 
than  does  the  Recent  species  ( Fig.  31 ) . 
Unless  the  differences  in  vertebral  meris- 
tics  are  also  considered,  however,  the 
morphometric  data  for  this  feature  are 
not  useful  in  comparing  the  various 
fossil  forms.  A.  uintaensis  and,  to  a 
lesser  extent,  A.  scutata  have  relatively 
longer  heads  with  a  vertebral  column  of 
approximately  85  centra.  A.  fragosa,  on 
the  other  hand,  is  a  short-bodied  form 
(approximately  65  centra)  with  a  short 
head.  A.  calva  has  a  relatively  long  ver- 
tebral column  (81-90  centra)  with  a 
shorter  head  than  the  other  long-bodied 
forms  (A.  uintaensis  and  A.  scutata). 

2.  Pelvic  fin  insertion  has  no  taxonomic  sig- 
nificance. 

3.  Anal  fin  insertion  may  have  minor 
ta.xonomic  significance  for  the  North 
American  specimens  of  A.  fragosa  wliich 
are  relatively  shorter  in  this  dimension 
than  in  the  other  .species.  There  is  too 
much    morphological    overlap   between 


18         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


the   species,   however,   to   make   this    a 
useful  criterion. 

4.  A.  uintaensis  has  a  relatively  longer 
mandible/head  ratio  (0.693)  than  any 
of  the  other  species  of  Amia,  while  A. 
frafiosa  has  a  smaller  ratio  (0.507 
mean).  The  mandible/head  ratio  of  A. 
scutata  (0.604  mean)  is  close  to  that  of 
A.  calva  ( 0.609  mean ) . 

5.  There  is  a  trend  from  the  Late  Creta- 
ceous to  Late  Miocene  in  the  lengthen- 
ing of  the  parietals  in  relation  to  the 
frontals.  Although  it  is  possible  to  dis- 
cern groups  that  fall  into  categories  of 
smaller  and  larger  ratios  (Table  7),  in- 
terspecific moi-phological  overlap  makes 
it  difficult  to  separate  any  one  of  the 
fossil  species  from  the  others  on  this 
criterion. 

6.  All  the  fossil  forms  have  operculum 
depth /width  proportions  that  fall  into 
the  lower  limits  of  the  A.  calva  range 
(0.921-1.025).  These  ratios  show  a 
slight  temporal  trend  towards  increasing 
width,  but,  while  this  is  perceptible,  it 
is  insufficient  to  indicate  taxonomic  sig- 
nificance. 

These  trends  suggest  possible  phylo- 
genetic  relationships  between  the  various 
amiid  species.  The  moiphometric  similari- 
ties indicate  that  little  evidence  exists  for 
maintaining  A.  scutata  and  A.  "dictijoce- 
phala"  as  separate  species.  The  Oligocene 
A.  scutata  is  distinguishable  quantitatively 
from  A.  calva  only  on  the  basis  of  a  larger 
head/ standard-length  ratio,  and  in  this  fea- 
ture it  is  intermediate  between  A.  calva  and 
A.  uintaensis.  The  moi-phometric  evidence 
indicates  similarities  between  A.  fra^,osa 
(Cretaceous-Eocene),  "Paramiatus  ii^iirleiji" 
( Eocene ) ,  and  A.  kehreri  ( Eocene ) .  Head/ 
standard-length  ratio  is  approximately  the 
same  among  these  three  forms;  insertion  of 
anal  and  pelvic  fins /standard-length  ratios 
shows  only  minor  differences.  Mandible/ 
head  size  and  parietal /frontal  ratios  are  al- 
most identical.  Of  all  the  .species,  A. 
uintaensis  is  the  most  morphometrically  dis- 
tinct.   It  has  a  relatively  greater  mandible/ 


head  ratio  and  a  smaller  parietal /frontal 
ratio  than  A.  fra^osa  (Table  7).  Even 
though  it  possesses  approximately  the  same 
total  number  of  centia  as  A.  calva  and  A. 
scutata  (Table  9),  it  still  has  a  greater 
head/ standard-length  ratio  than  the  two 
latter  species.  Temporally,  there  are  minor 
trends  in  Amia  towards  lengthening  of  the 
parietals  in  relation  to  the  frontals,  and  in- 
creasing operculum  width  to  depth. 

MERISTICS 

Meristic  elements  have  been  used  in 
species  diagnoses  of  various  fossil  amiids 
by  Cope  (1875),  Osborn  et  al.  (1878), 
Romer  and  Fryxell  (1928),  and  Estes 
( 1964 ) .  A  meristic  study  of  both  Recent 
and  fossil  species  of  Amia  was  undertaken 
to  determine  tlie  relative  value  of  such 
diagnoses  in  the  taxonomy  of  the  amiids. 
A  comparison  of  the  number  of  supraverte- 
bral  scale  rows,  the  number  of  branchio- 
stegal  rays,  and  the  number  of  pectoral, 
pelvic,  anal,  dorsal,  and  caudal  fin  rays  com- 
prises the  first  part  of  the  study,  while  com- 
parative vertebral  meristics  comprise  the 
second  part. 

Supravertebral  Scale  Rows 

Cope  (1875)  differentiated  A.  scutata  from 
A.  calva  and  A.  "dictyocephala"  on  the  basis 
of  A.  scutata  s  (USNM  5374)  having  seven 
and  a  half  longitudinal  rows  of  large  scales 
above  the  vertebral  column.  Cope  (1875) 
described  A.  "dictyocephala"  (USNM  3992) 
as  bearing  ten  to  twelve  rows  of  scales 
above  the  vertebral  column.  A  count  of  the 
scale  rows  between  dorsal  fin  distal  pterygi- 
ophores  and  the  vertebral  column  in  20  Re- 
cent A.  calva  (Table  8)  gave  a  range  of 
seven  to  nine  supravertebral  scale  rows. 
Although  the  number  of  scale  rows  will 
vary  with  the  region  of  the  trunk  anatomy 
from  which  the  count  might  be  taken,  Cope 
did  not  designate  the  point  at  which  he 
made  his  scale  row  count.  Also,  his  speci- 
men WAS  so  poorly  preserved  that  his  count 
may  have  been  affected  by  distortion  of 
the  scales.  The  only  way  that  a  valid  com- 
parison of  all  the  fonns  could  be  made  was 


Fossil  Amiius  •  Boreskc 


19 


TaULK    fS.        (loMl'AHlSON   OF   MKHISTIC:   ELEMENTS   IN    HECENT  AND  FOSSIL   NOHTH  AMERICAN   AMIIDS 


Supra- 

Branchi- 

Pecf  oral-Fin 

Pelvic-Fin 

Anal-Fin 

Dorsal-Fin 

C"aiidal-Fin 

vertebral 

ostegal 

Lepido- 

Lepido- 

I.epido- 

Lepido- 

Lcpido- 

Scale  Rows 

Rays 

trichia 

trichia 

Irichia 

trichia 

tridiia 

Amia  calva  (20) 

7-9 

10-13 

16-19 

7-8 

8-11 

45-49 

23-27 

Recent 

7.5  av. 

11.4  av. 

16.8  av. 

7.2  av. 

10.5  av. 

48.0  av. 

25.7  av. 

A.  sciitata 

YPM  6243" 

USNM  4087* 

PU  10172' 

7 

11 

— 

7 

9 

47""' 

A.  scutata 

USNM  5374 

7.5 

— 

— 

9 

— 

A.  scutata 

YPM  6241 

8 

9 

— 

23 

A.  scutata 

UMMP  V-57431 

7 

11 

17 

7 

9 

46** 

A.  "dictijoccphala" 

USNM  3992 

7.8 

— 

— 

7 

9 

48»» 

— 

A.  "dicttjoccphaJa" 

AMNH  2802 

11 

— 

— 

A.  "dictijoccphala" 

AMNH  2670 

9*' 

— 

— 

470  » 

A.  iiintacnsis 

PU  13865 

7 

16 

9 

9"* 

23 

A.  uintacnsis 

AMNH  785 

9«o 

— 

7 

10 

— 

24 

A.  fragosa 

MCZ  5341 

8 

12 

18 

7 

8 

45 

19-20 

"Paramiatus  gu dcyi" 

FMNH  2201 

7-8 

12 

17 

8 

8 

44-45** 

19 

A.  fragosa 

UA  5506 

10 

— 

A.  fragosa 

UA  5425 

— 

19 

"  All  one  specimen. 
"o  Est. 


to  take  the  supravertebral  scale  row  count 
of  both  the  USNM  3992  specimen  and  the 
other  fossil  and  Recent  amiid  specimens  at 
the  same  point.  In  this  case,  I  took  all 
connts  on  a  vertical  line  at  the  level  of  the 
posterior  pterygiophore  of  tlie  anal  fin.  I 
connted  the  nnmber  of  scale  rows  in  speci- 
mens of  A.  frcif^osa,  A.  uintaensis,  and  "Para- 
miatus ^urleyi"  in  addition  to  those  of 
Cope's  types  of  A.  scutata  (USNM  5374) 
and  A.  "dictyocephala"  (USNM  3992),  as 
well  as  referred  specimens  of  A.  scutata; 
I  then  compared  them  with  the  supraverte- 
bral scale  row  range  in  A.  calva.  The 
supravertebral  scale  rows  of  fossil  Amia 
(Table  8)  appear  to  fall  within  the  supra- 
vertebral scale  row  range  of  Recent  A. 
calva.    Although   Cope   had   described   A. 


"dictyocephakr  (on  the  basis  of  USNM 
3992)  as  having  10-12  scale  rows,  I  believe 
his  count  is  too  high.  The  supravertebral 
scale  rows  in  this  and  other  fossil  forms  are 
difficult  to  observe  for  several  reasons. 
Amiid  scales  are  aligned  diagonally  to  the 
vertebral  column  rather  than  in  parallel, 
making  it  often  difficult,  particularly  in 
fossil  material,  to  determine  to  which  diag- 
onal column  the  overlapping  scale  rows 
belong.  Also,  the  scales  on  the  USNM  3992 
specimen  are  broken  into  many  parts,  and 
Cope  may  therefore  have  been  counting 
partial  scales  as  whole  ones.  I  believe  that 
I  obtained  a  more  reasonable  estimate  of 
the  supravertebral  scale  row  count  in  this 
specimen  in  the  following  manner:  I 
measured  the  average  of  the  anteroventral 


20         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


width  of  complete  scales  from  the  abdomi- 
nal region  (in  which  the  scales  are  the 
same  size  as  in  other  places  in  the  mid-body 
region)  and  then  divided  that  amount  into 
the  distance  between  the  midpoint  of  the 
vertebral  column  and  the  dorsal  fin  distal 
pterygiophore.  In  this  case,  the  quotient 
was  7.8,  which  is  comparable  with  the 
counts  of  approximately  7-9  in  the  other 
Oligocene  specimens  and  in  A.  calva  ( Table 
8).  No  taxonomic  significance  can  thus  be 
applied  to  the  number  of  scale  rows  above 
the  vertebral  column  since  counts  in  Recent 
and  fossil  Amia  fall  within  a'  relatively 
narrow  range. 

Branchiostegal  Rays 

The  number  of  branchiostegal  rays  was 
included  in  the  species  diagnosis  for  A. 
"dictyocephalo"  (AMNH  2802),  in  which 
Cope  ( 1875)  counted  12  rays.  Osborn  et  al. 
(1878)  observed  13  branchiostegal  rays  in 
A.  scutata  (PU  10172)  and  Romer  and 
Fryxell  (1928)  figured  12  such  rays  for 
"Paramiatus  ^urleyi"  (FMNH  2201). 
O'Brien  (1969)  counted  10  rays  in  A. 
fragosa  (UA  5506)  from  the  Edmonton 
Formation.  On  the  basis  of  disarticulated 
material  from  the  Late  Cretaceous  Lance 
Formation,  Estes  (1964)  estimated  that  A. 
fragosa  would  bear  14  branchiostegal  rays, 
like  the  Late  Jurassic  Sinamia  zdanskyi  de- 
scribed from  China  by  Stensio  (1935;  see 
Liu  et  al,  1963  for  range  and  distribution). 
In  the  Recent  A.  calva  sample  (Table  8), 
the  number  of  branchiostegal  rays  ranges 
from  10  to  13;  the  range  among  the  few 
known  examples  of  fossil  forms  is  from  10 
to  13,  an  indication  that  the  number  of 
branchiostegal  rays  has  remained  constant. 

Fin  Rays 

Because  of  confusing  duplication  of  ter- 
minology used  for  fin  description  in  the 
literature,  I  will  use  that  of  Lagler  et  al. 
( 1962 )  for  the  appendicular  skeleton  unless 
I  indicate  otherwise. 

All  fin  ray  counts  on  Recent  A.  calva 
were  obtained  from  X-rays  of  20  specimens 
from  Wisconsin  and  Michigan.   The  counts 


taken  from  fossil  forms  are  as  accurate  as 
conditions  allow,  although  a  number  of  the 
specimens  are  incomplete  or  show  only 
traces  of  the  actual  fin.  The  results  of  this 
study  must  therefore  be  considered  with 
this  in  mind.  I  ol^tained  these  counts  from 
as  close  as  possible  to  the  internal  fin  sup- 
ports rather  than  to  the  segmented  and  bi- 
furcated distal  lepidotrichia.  There  is  a 
one-to-one  correspondence  between  the 
number  of  lepidotrichia  and  the  number  of 
pterygiophores  in  the  anal  and  dorsal  fins; 
however,  this  is  not  the  case  in  the  pectoral, 
pelvic,  and  caudal  fins,  which  have  more 
lepidotrichia  than  fin  supports  (Fig.  31). 

Pectoral  fin.  The  number  of  pectoral  fin 
lepidotrichia  has  not  been  previously  noted 
in  any  of  the  original  species  descriptions 
of  fossil  Amia.  There  are  four  .specimens  in 
which  it  is  possible  to  make  a  pectoral  fin 
ray  count  (Table  8).  A.  scutata  (1),  A. 
uintaensis  (1),  and  A.  fragosa  (2)  speci- 
mens bear  16  to  18  pectoral  lepidotrichia,  a 
number  which  is  approximately  the  average 
for  20  specimens  of  Recent  A.  calva  which 
displayed  from  16  to  19  pectoral  fin  rays 
(Table  8).  O'Brien's  (1969)  analysis  of  A. 
fragosa  (Edmonton  Formation)  does  not 
include  any  quantitative  comparison  of  its 
pectoral  fins  with  those  of  A.  calva.  He 
does,  however,  observe  that  the  pectoral 
fins  are  qualitatively  similar  in  the  two 
species.  The  pectoral  fins  of  A.  fragosa,  A. 
scutata,  and  A.  uintaensis  thus  do  not  vary 
meristically  from  those  of  A.  calva.  Lehman 
(1951:  8),  in  his  description  of  Pseudamia 
heintzi  from  the  Eocene  of  Spitzbergen, 
notes  that  the  pectoral  fin  has  13  complete 
nonbifurcating  lepidotrichia  in  the  visible 
portion  of  the  fossil.  This  count  is  different 
from  that  of  both  Recent  and  fossil  North 
American  amiids,  but  as  Lehman's  plate  3 
indicates,  this  difference  may  be  caused  by 
matrix  that  overlies  the  ventral  portion  of 
the  pectoral  fin,  possibly  covering  addi- 
tional lepidotrichia. 

Pelvic  fin.  The  number  of  lepidotrichia 
of  the  pelvic  fin  was  part  of  Cope's  ( 1875 ) 
species  diagnosis  for  A.  "dictyocephala" 
(USNM  3992)   and  that  of  Osborn  et  al. 


Fossil  Amiids  •  Boreske        21 


(1878)  for  A.  scutota  (PU  10172).  I 
counted  the  lepidotrichia  of  these  speci- 
mens as  well  as  those  of  on(>  additional 
Ohgocene  specimen  and  compared  them 
with  my  sample  of  A.  calva,  which  showed 
between  seven  and  eight  pehic  lepido- 
trichia (Table  8).  Although  Osborn  et  al. 
( 1878)  counted  ten  pelvic  lepidotrichia,  my 
recount  of  their  A.  scutata  specimen  (PU 
10172)  showed  only  seven  (Plate  4).  The 
bifurcation  of  the  fin  rays  might  have  been 
inadvertently  included  in  their  original 
count.  The  holotype  of  A.  "dictyocepliala" 
(USNM  3992)  (Fig.  27)  showed  seven 
rather  than  the  six  lepidotrichia  that  Cope 
(1875)  had  diagnosed.  A  specimen  of  A. 
scutata  (UMMP  V-57431)  (Fig.  27A)  also 
has  seven  lepidotrichia;  both  of  these  are 
within  the  range  of  Recent  A.  calva.  Of  the 
remaining  fossil  forms,  A.  fraii^osa  and 
"Paramiatus  ^urleyi"  have  eight,  and  A. 
uintaensis  nine,  A.  uintacnsis  being  the  only 
fossil  form  not  to  fall  within  the  range  of 
Recent  A.  calva.  This  difference  is  insuf- 
ficient to  demonstrate  any  taxonomic  value, 
however,  at  least  until  more  A.  uintaensis 
specimens  are  known. 

Anal  fin.  Anal  fin  lepidotrichia  have 
been  included  in  the  diagnoses  of  A.  "dic- 
tyocephala"  and  A.  scutata  (Cope,  1875), 
and  also  in  the  description  of  A.  .scutata 
(Osborn  et  al,  1878).  Each  of  the  original 
counts  of  nine  anal  rays  for  each  specimen 
concurs  with  my  recount  and  also  falls 
within  the  range  of  eight  to  eleven  for  Re- 
cent A.  calva  (Table  8).  A.  jra<i,osa,  "Para- 
miatus pMrleyi"  and  A.  uintaensis  also  fall 
within  the  range  of  A.  calva. 

Dorsal  fin.  Although  the  number  of 
lepidotrichia  in  the  dorsal  fin  has  been 
mentioned  by  several  authors  in  their  diag- 
noses of  fossil  amiids,  it  is  one  of  the  more 
difficult  meristic  counts  to  obtain,  since  a 
complete  dorsal  region  of  the  fossil  is 
required.  Cope's  type  of  A.  "dictyocephala" 
(USNM  3992)  lacks  a  complete' dorsal  fin, 
so  he  counted  only  the  32  dorsal  lepidotri- 
chia between  the  beginning  of  the  dorsal 
fin  and  the  posterior  lepidotrichia  of  the 
anal    fin    (Cope,    1875).     Osborn    et    al. 


(1878)  reported  53  dorsal  lepidotrichia  for 
A.  scutata  (PU  10172),  but  this  must  have 
been  an  estimate,  since  the  posterior  por- 
tion ot  the  dorsal  fin  as  well  as  the  entire 
caudal  fin  is  missing  (Plate  4C).  As  the 
two  A.  scutata  specimens  with  complete 
dorsal  fins  (AMNM  2670,  UMMP  V- 
57431)  have,  respectively,  47  and  46  dorsal 
lepidotrichia  (Table  8),  it  seems  that  the 
count  of  Osborn  ct  al.  ( 1878)  was  high  and 
that  the  PU  10172  specimen  would  prob- 
ably have  corresponded  with  the  other 
Oligocene  specimens. 

Romer  and  Fryxell's  ( 1928 )  diagnosis  for 
"Paramiatus  fiurleyi"  includes  a  dorsal  fin 
ray  count  of  45,  which  they  note  as  being 
slightly  fewer  than  the  count  for  A.  calva. 
O'Brien's  ( 1969 )  discussion  of  Edmonton 
Formation  A.  fra^osa  does  not  include  any 
counts  of  dorsal  lepidotrichia,  although  he 
does  note  that  the  relative  length  of  the 
entire  dorsal  fin  in  A.  fraiiosa  is  similar  to 
that  of  A.  calva.  In  the  A.  calva  specimens 
I  studied,  the  dorsal  fin  rays  ranged  be- 
tween 45  and  49,  the  average  approximately 
48.  Romer  and  Fryxell's  diagnosis  of  "Para- 
miatus fiurleyi"  as  having  slightly  fewer 
dorsal  lepidotrichia  than  A.  calva  is  correct, 
but  this  and  all  the  related  fossil  forms  fall 
within  the  lower  range  of  A.  calva  (Table 
8).  The  number  of  dorsal  fin  rays  appears 
to  have  little  taxonomic  value. 

It  is  interesting  that  the  complete  Amia 
frafi:osa  (MCZ  5341),  "Paramiatus  <^urleyi" 
(FMNH  2201),  and  A.  kehreri  (BMNH 
P33480)  specimens  have  dorsal  fins  of 
nearly  the  same  length  and  contain  the 
same  number  of  lepidotrichia  as  A.  calva, 
despite  the  fact  that,  on  the  basis  of  the 
number  of  vertebrae,  these  species  have  a 
much  shorter  body  (Table  9).  This  con- 
tributes to  a  proportional  difference  in  the 
body  forms  of  these  species,  since  the  dorsal 
fin  in  A.  fra<^o.sa  (including  "Paramiatus 
il,urleyi")  terminates  much  closer  to  the 
caudal  fin  than  in  A.  calva  (Plate  1;  Fig. 
31).  However,  as  Shufeldt  (1885)  and  Hay 
(1895)  implied,  it  is  very  doubtful  that  the 
dorsal  fin  was  fused  into  a  continuous  struc- 
ture with  the  caudal  fin  in  some  ancestral 


22         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


amiid.  The  Late  Mesozoic  European  forms 
of  Urocles  and  Amiopsis  have  a  much  ab- 
breviated dorsal  fin  that  terminates  more 
anteriorly  than  does  that  of  the  species  of 
Amia.  For  Amiopsis  dolloi,  an  Early  Creta- 
ceous (Wealden)  amiid  from  Bernissart, 
Belgium,  Traquair  (1911)  figured  17  dorsal 
fin  supports,  while  Lange  (1968)  estab- 
lishes a  specific  range  of  17-25  for  the  Eu- 
European  Upper  Jurassic  Urocles.  The 
basis  of  Shufeldt's  (1885:  8.5-86)  model  for 
a  primitive  amiid  with  a  continuous  dorsal- 
caudal  fin  was  the  presence  in  Recent  Amia 
calva  specimens  of  what  Shufeldt  called  a 
"series  of  delicate  little  bones  that  continue 
the  interspinous  bones  of  the  dorsal  fin  as 
far  as  the  caudal  fin."  These  five  bones  he 
considered  to  be  the  continuation  of  the 
dorsal  interneural  spines;  Hay  (1895),  in 
his  discussion  of  Amia  calva,  refers  to  them 
as  "epural  interspinous  bones."  These  small 
bones  can  also  be  seen  in  several  of  the 
fossil  amiids  I  have  studied,  especially  A. 
scutata  (YPM  6241),  A.  fragosa  (UA  5425), 
and  A.  uintaensis  (AMNH  785)  (Fig.  8). 
From  these  fossil  forms,  however,  it  is  dif- 
cult  to  determine  whether  the  origin  of 
these  bones  is  from  the  dorsal  or  caudal 
neural  spines.  An  examination  of  the  caudal 
fin  of  Urocles  lepidotus  ( Fig.  6;  also 
Nybelin,  1963:  506,  fig.  17),  which  is 
known  to  have  an  abbreviated,  more  an- 
teriorly located  dorsal  fin,  shows  that  these 
epural  bones  are  associated  with  the  caudal 


fin,  which  supports  the  upper  caudal  lepi- 
dotrichia  in  much  the  same  manner  as  the 
hypurals  in  the  ventral  tail  region.  A 
further  indication  that  these  epural  inter- 
spinous bones  are  not  vestigial  dorsal  spine 
supports  is  found  in  Traquair's  ( 1911 )  plate 
7  of  Amiopsis  dolloi  and  his  plate  8  of 
Amiopsis  lata  (both  species  from  the  Creta- 
ceous [Wealden]  of  Belgium);  these  plates 
show  the  bones  to  be  clearly  associated 
with    the    caudal    lepidotrichia    (Fig.    7). 


i 


Fig.  6.      Urocles  lepidofus  MCZ  8300,  caudal  fin. 


Fig.  7.  Restoration  of  Amiopsis  dolloi,  scales  omitted 
Early  Cretaceous  (Wealden),  Bernissart,  Belgium  (after 
Traquair,  1911). 


Fossil  Amiids  •  Boreske        23 


There  are  no  intermediate  interspinous 
bones  between  these  bones  in  the  caudal 
region  and  those  of  the  much  more  anteri- 
orly situated  dorsal  spine.  The  fin  of  a  648- 
mm  SL  Recent  Amia  calva  (Fig.  10)  does, 
however,  confirm  tliat  Shuf eldt  ( 1885 )  was 
correct  in  stating  that  the  epurals  are  con- 
tinuations of  the  interneural  spines.  Figure 
10  shows  three  free  interspinous  epurals, 
with  a  fourth  that  is  either  being  fused  onto 
a  neurd  spine  or  is  actually  a  single  greatly 
elongated  neural  spine.  As  only  two  of 
these  epurals  are  attached  to  lepidotrichia, 
there  is  not  a  one-to-one  correspondence 
between  the  two  elements,  as  in  the  hy- 
purals  in  the  main  caudal  region. 

Caudal  fin.  With  the  exception  of 
Romer  and  Fryxell's  (1928)  diagnosis  of 
"Paramiatus  g,urleyi,"  none  of  the  original 
descriptions  of  fossil  Amia  include  counts  of 
the  caudal  fin  rays.  Although  Romer  and 
Fryxell  observed  20  caudal  lepidotrichia,  a 
recount  shows  only  19  (Fig.  8E).  Other 
fossil  forms  tliat  also  show  19  caudal  fin 
rays  are  A.  fragosa  (UA  5425)  from  the 
Edmonton  Formation,  A.  fragosa  (MCZ 
5341 )  from  the  Green  River  Formation, 
and  A.  kehreri  from  Messel  (Andreae, 
1895,  plate  1,  fig.  23).  Another  specimen 
of  A.  kehreri  from  Messel  (BMNH  P33480) 
has  18  lepidotrichia  (Plate  2).  Traquair's 
( 1911 )  plate  7  of  three  specimens  of  Amiop- 
sis  dolloi  shows  between  15  and  17  caudal 
lepidotrichia,  while  Urocles  spp.  have  a 
range  between  12-18  caudal  lepidotrichia 
(Lange,  1968).  The  only  Oligocene  speci- 
men with  a  complete  caudal  fin  (YPM 
6241)  has  23  caudal  lepidotrichia;  the 
Eocene  specimens  of  Amia  uintaensis  show 
23  to  24.  Although  my  sample  of  Amia 
calva  displays  caudal  fins  with  a  range  of 
23  to  27  lepidotrichia,  the  number  of  these 
caudal  fin  rays  is  skewed  toward  the  higher 
limit  of  the  range  (Fig.  9).  There  is  thus 
a  considerable  difference  between  the  num- 
ber of  lepidotrichia  in  Amia  fragosa  and 
the  majority  of  the  A.  calva  specimens.  A. 
scutata  is,  however,  within  the  range  of  the 
Recent  species,  but  occupies  the  lower 
limits  of  the  range. 


Thus,  of  all  the  meristic  elements  so  far 
considered,  it  appears  that  the  greatest  dis- 
parity between  the  fossil  forms  and  the 
Recent  A.  calva  is  in  the  number  of  caudal 
fin  rays.  The  number  of  caudal  fin  rays 
therefore  appears  to  have  taxonomic  impor- 
tance and  may  have  some  functional  as  well 
as  morphological  correspondence  to  the  two 
different  amiid  body  types. 

As  discussed  in  the  preceding  section  on 
dorsal  fins,  there  are  two  attachment  bases 
for  the  caudal  lepidotrichia:  epural  inter- 
spinous bones  and  the  hypurals.  The 
epurals  are  usually  attached  to  only  two  or 
three  of  the  caudal  fin  rays,  while  the 
remainder  of  the  lepidotrichia  are  sup- 
ported by  the  hypurals.  Nybelin  {in  1963: 
488)  defines  hypurals  as  "those  haemal  ele- 
ments located  to  the  rear  of  the  emergence 
of  the  caudal  artery  from  the  haemal  canal" 
(trans.  Lund,  1967:  210)  (Fig.  lOB).  Lund 
(1967:  210)  agrees  instead  with  White- 
house  (1910:  592),  who  defines  hypurals 
as  "any  hypaxial  elements  that  support 
caudal  fin  rays"  (Fig.  lOA).  Lund  states 
that  the  sole  function  of  a  hypural  is  to 
support  a  caudal  fin  ray  and  therefore  the 
first  hypural  would  be  "the  first  haemal 
spine  in  rem'ward  progression  to  support  a 
caudal  fin  ray  and  the  first  ural  centrum 
is  the  centrum  supporting  the  first  hypural 
element."  Lund's  definition  is  more  practical 
for  paleontological  use.  Since  there  is  an 
intennediate  joint  (Figs.  8,  10).  the  major- 
ity of  the  hypurals  are  not  attached  directly 
to  the  urals.  However,  as  Shufeldt  (1885) 
and  Hay  ( 1895 )  observed,  the  posterior- 
most  seven  to  nine  hypurals  are  ankylosed 
to  the  corresponding  vertebrae  ( Fig.  IOC ) . 
This  same  co-ossified  condition  of  the  last 
hypurals  is  also  evident  in  the  fossil  fonns, 
so  that  the  number  of  these  fused  hypurals 
has  remained  constant  throughout  the  evo- 
lutionary history  of  Amia.  Also,  as  Figure 
10  shows,  the  seven  or  eight  anteriormost 
hypurals  of  Recent  A.  calva  have  a  one-to- 
two  correspondence  with  the  ventral  lepi- 
dotrichia. In  most  of  the  available  fossil 
amiid  specimens,  the  ventral  caudal  portion 
is  poorly  preserved,  so  that  it  is  difficult  to 


24         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Fig.  8.     Caudal    regions:    A,    Amia    scufaia    YPM    6241;    B,    A.    scufafa    AMNH    2671;    C,    A.    uintaensis    AMNH    785; 
D,  A.  fragosa  UA  5425;  and  E,  A.  fragosa  FMNH  2201. 


Fossil  Amiids  •  Boreske        25 


u 

a> 


11    ^n    ^li    ^5       2r~ 

Caudal  Fin  Lepidotrichia  no. 


TT 


Fig.  9.  Number  of  caudal  lepidotrichia  in  20  speci- 
mens of  Recent  Ami'a  calva. 

arrive  at  an  accurate  count  of  the  total 
number  of  hypurals  or  to  verify  whether 
this  one-to-tvvo  relationship  exists  in  all  the 
amiid  fossil  forms.  The  only  available  fossil 
form  in  which  this  one-to-two  hypural- 
lepidotrichia  coiTCspondence  in  the  ventral 
caudal  region  can  clearly  be  seen  is  in  A. 
scutata  (YPM  6241;  Fig.  8). 

Vertebral  Elements 

Two  regions  of  the  vertebral  column,  the 
trunk  and  the  caudal  regions,  are  defined  by 
their  relationships  to  the  ribs,  neural  arches, 
and  haemal  arches.  The  trunk  region  con- 
sists of  monospondylous  vertebrae  that  pos- 
sess paired  basapophyses  having  gradually 
changing  angles,  dorsal  neural  facets,  and 
ventral  aortal  facets.  The  number  of  tnmk 
vertebrae  in  my  sample  of  Ainia  calva  varies 
from  36  to  38.  The  caudal  region  consists 
of  three  types  of  vertebrae,  listed  from  an- 
terior to  posterior:  regular  monospondylous 
centra  bearing  neural  and  haemal  arches, 
diplospondylous  centra  bearing  neither 
neural  nor  haemal  arches  (neural  and 
haemal  facets  still  present ) ,  and  ural  centra. 
Since  the  neural  and  haemal  facets  are  still 
present  in  the  diplospondylous  centra,  there 


is  no  way  to  differentiate  the  latter  from 
the  monospondylous  type  in  a  disarticulated 
state.  In  my  sample  of  A.  calva,  the  number 
of  regular  caudal  monospondylous  centra 
(24-26)  fluctuates  by  two  centra,  that  of 
the  diplospondylous  caudal  centra  ( 14-17 ) 
by  three  (Table  9). 

The  posterior  caudal  region  of  A.  calva 
consists  of  two  types  of  urals:  centra  with 
hypurals  attached  by  a  layer  of  cartilage 
(free  urals),  and  centra  that  are  fused 
directly  onto  the  hypurals,  often  lacking  the 
neural  arches  (fused  urals).  When  dis- 
articulated, the  fused  urals  can  often  be 
distinguished  from  the  free  urals,  since 
part  of  the  hypural  usually  remains  fused 
to  the  ural,  extending  the  posterior  articular 
surface  downward.  The  nonfused  (free) 
urals  cannot  be  distinguished  in  a  disarticu- 
lated state  from  the  monospondylous  or 
diplospondylous  caudal  centra.  The  num- 
ber of  urals  with  fused  hypurals  is  readily 
counted,  since  they  are  distinguishable 
from  the  remainder  of  the  vertebrae.  In 
order  to  identify  a  free  ural,  it  is  necessary 
to  observe  the  relationship  between  the 
ural  and  its  conesponding  hypural  and 
lepidotrichia.  It  is  often  difficult  to  make 
this  distinction  between  free  and  fused 
urals,  since  the  caudal  region  is  seldom 
complete  in  articulated  fossil  forms.  In  A. 
calva  the  number  of  urals  with  ankylosed 
hypurals  ranges  between  seven  and  nine. 
There  are  approximately  seven  principal 
urals  fused  to  hypurals,  followed  by  one  or 
two  small  additional  urals  that  do  not  ar- 
ticulate with  the  preceding  vertebrae  but 
lie  dorsal  to  the  upturned  portion  of  the 
vertebral  column.  Because  it  is  difficult  to 
discern  these  urals  in  smaller  specimens  of 
A.  calva,  the  count  may  be  slightly  biased, 
and  a  comparison  of  the  fossil  forms  with 
the  range  established  for  A.  calva  must  be 
made  with  this  consideration  in  mind. 

I  counted  the  number  of  centra  between 
the  anterior  dorsal  fin  pterygiophore  and 
the  posterior  anal  fin  pterygiophore,  since 
Cope  (1875)  used  the  number  of  central 
elements  between  these  points  as  a  specific 
character  for  A.   "dictyocephala"   (USNM 


26         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Fig.    10.     Am'ia    calva    (648    mm    SL)    caudal:    A,    Whitehouse    (1910)    and    Lund's    (1967)    definition    of    first    ural;    B, 
Nybelin's  (1963)  definition  of  first  ural;  C,  first  fused  ural. 


3992 ) .  The  range  for  the  number  of  centra 
in  this  region  of  Recent  A.  calva  is  33  to  37. 
There  is  considerable  variation  in  total 
number  of  centi'a  {i.e.,  segments)  in  Recent 
A.  calva  (81-90),  which  may  pose  a  prob- 
lem in  comparing  specific  vertebrae.  Thus 
in  two  A.  calva,  for  example,  the  eightieth 
vertebral  segment  of  one  individual  might 
not  correspond  to  the  same  position  in  the 
vertebral  column  or  even  type  of  centrum 
as  the  eightieth  segment  of  the  second  indi- 
vidual. This  should  be  considered  in  any 
comparisons  of  several  A.  calva  individuals, 
as  well  as  in  comparisons  of  the  fossil  forms, 
which  share  this  variation  in  vertebral  seg- 
ments (Table  9).  Also,  fusion  of  vertebral 
elements  may  occur  in  Recent  A.  calva.  In 
some  specimens,  as  many  as  five  centra 
were  found  fused  together  at  points 
throughout  the  vertebral  column;  this  con- 
dition was  present  to  a  lesser  degree  or 
absent  in  other  specimens  (Tables  10-12). 
These  fused  centra  also  occur  in  the  fossil 
forms,  as  in  A.  uintaensis  (YPM  6244).  The 


actual  number  of  such  fused  centra  can 
often  be  established  only  by  counting  ex- 
ternal features  such  as  basapophyses,  neural 
facets,  aortal  facets,  or  haemal  facets. 

Romer  and  Fryxell's  ( 1928 )  study  of 
"Paramiatus  gurleyi"  is  the  only  pubHshed 
description  of  a  complete  articulated  fossil 
amiid.  They  distinguished  this  form  from 
the  Recent  species  by  the  supposed  pres- 
ence of  a  deeper  body,  and  also  noted  that 
the  number  of  centra  was  considerably  less 
than  in  A.  calva.  The  vertebral  column  is 
completely  preserved,  so  that  it  is  possible 
to  obtain  an  accurate  count  of  the  vertebrae 
(Plate  IB).  "Paramiatus  p,iirleyi"  has  67 
vertebral  segments  in  contrast  to  the  mean 
of  86  in  A.  calva  (Table  9).  Osborn  et  al. 
(1878)  described  A.  sciitata  (PU  10172)  on 
the  basis  of  a  specimen  lacking  a  caudal 
fin  (Plate  4).  Since  the  specimen  is  other- 
wise complete,  they  were  able  to  estimate 
that  their  specimen  had  82  vertebral  seg- 
ments. 

Cope    (1875)    described    A.    "clictyoce- 


Fossil  Amiids  •  Boreske        27 


Table  9. 

Comparison  of 

VERTEBRAL 

CHARACTERS    IN 

RECENT   AND 

FOSSIL   AMUDS 

Number  of 

Centra 

between 

Anterior 

Number 

Number 

Number 

Dorsal-Fin 

of  Mono- 

of  Diplo- 

of  Ural 

Pterygiophore 

Total 

Number  of 

spondylous 

spondylous 

Centra 

and  Posterior 

Number  of 

Trunk 

Caudal 

Caudal 

with  Fused 

Anal-Fin 

Centra"" 

Centra 

Centra 

Centra 

Hypurals 

Pterygiophore" 

Recent 

Amia  calva  (20) 

Wis.  &  Mich. 

81-90 

36-38 

24-26 

14-17 

7-9 

33-37 

mean 

mean 

mean 

mean 

mean 

mean 

=  85.8 

=  37.3 

=  25.2 

=  16.2 

=  8.3 

=  35.5 

Oligocene 

A.  scutata 

PU  10172 

83*** 

36 

25 

15 

yooo 

35 

A.  scutata 

UMMP  V-57431 

81*** 

36 

24*** 

15*** 

Y*  00 

37 

A.  "dicttjoccphala" 

USNM  3992* 

— 

— 

35 

Eocene 

"Paramiatus  ^urlcyi 

•* 

FMNH  2201* 

67 

26 

19 

16 

6 

26 

Amia  uintaensis 

PU  13865 

85 

31 

26 

21 

7 

36 

Amia  uintaensis 

AMNH  785 

25 

20 

7 

— 

A.  fragosa 

MCZ  5341 

65 

25 

18 

15 

7 

25 

A.  kehreri 

BMNH  P33480 

62*** 

24 

16 

16 

6*** 

24 

types. 


o   -- 

""  ^  including  diplospondylous  units    (as  one), 
"o"  Est. 


pluild"  from  a  specimen  (USNM  3992)  in 
which  only  the  mid-body  region  was  pre- 
served. He  felt  that  the  number  of  ver- 
tebrae between  the  anterior  dorsal  fin 
pterygiophore  and  the  posterior  anal  fin 
pterygiophore  had  ta.xonomic  significance. 
A  comparison  of  this  specimen  with  Recent 
A.  calva  showed  that  the  v'ertebral  count  of 
this  region  is  essentially  the  same  in  both 
species.  This  character  is  therefore  not 
useful  in  distinguishing  this  species  from 
the  Recent  form  or  in  characterizing  it  as 
a  specific  taxon.  The  specimens  of  A. 
scutata  are  within  the  range  of  A.  calva  in 
total  number  of  vertebrae  as  well  as  in  the 
number  of  vertebrae  in  the  vimous  cate- 
gories (Table  9) .  Based  on  the  similarity  of 
number  of  vertebrae  in  A.  scutata  to  that  of 
A.  calva,  it  appears  that  the  amiid  vertebral 


column  has  not  changed  meristically  from 
Oligocene  to  Recent. 

Additional  data  from  five  undescribed 
fossil  amiid  specimens  with  relatively  com- 
plete axial  skeletons  has  been  of  consider- 
able help  in  estimating  vertebral  counts 
of  the  fossil  forms.  A  complete  specimen 
of  A.  uintaensis  from  the  Green  River  For- 
mation (PU  13865)  has  a  complete  axial 
skeleton  (Plate  3).  Interestingly,  the  total 
number  of  centra  (85)  does  not  differ  from 
that  of  A.  scutata  or  A.  calva  (Table  9). 
The  only  variation  is  in  the  number  of 
trunk  centra  and  the  number  of  diplospon- 
dylous caudal  centra.  There  are  fewer 
trunk  centra  in  this  specimen  of  A.  uintaen- 
sis (31)  than  in  A.  scutata,  which  has  a 
mean  of  36,  or  in  A.  calva,  \\'hose  trunk 
centra  are  a  mean  of  37.    A  partially  com- 


28 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


plete  A.  uintaensis  specimen  (AMNH  785), 
also  from  the  Green  River  Formation,  shows 
almost  the  same  number  of  diplospondylous 
caudal  centra  as  PU  13865  (20-21  respec- 
tively). The  lesser  number  of  trunk  centra 
in  both  specimens  of  A.  uintaensis  is  thus 
offset  by  a  greater  number  of  diplospondy- 
lous caudal  centra.  In  comparing  the  verte- 
bral column  of  A.  uintaensis  with  that  of 
A.  calva,  A.  scutata,  and  A.  fragosa,  it  ap- 
pears that  although  A.  uintaensis  shares  the 
same  total  number  of  vertebral  segments 
with  A.  scutata  and  A.  calva,  it  does  not 
conform  to  their  proportional  division  of 
the  column  into  trunk  and  caudal  regions. 
A.  uintaensis  has  a  trunk/ total-number  ver- 
tebral ratio  of  0.365,  while  A.  fragosa  has  a 
ratio  of  0.300  as  compared  to  the  A.  calva 
ratio  of  0.440.  Three  complete  specimens 
referred  to  here  as  A.  fragosa  ("Paramiatus 
gurleyi"  FMNH  2022,  A.  kehreri  BMNH 
P33480,  and  A.  fragosa  MCZ  5341)  have 
vertebral  columns  that  differ  proportion- 
ately and  meristically  from  A.  calva,  A. 
scutata,  and  A.  uintaensis.  A.  fragosa  has 
significantly  fewer  centra  than  the  other 
fossil  forms,  with  approximately  12  fewer 
trunk  vertebrae  and  8  fewer  monospondy- 
lous  caudal  centra.  It  has  approximately 
the  same  number  of  diplospondylous  caudal 
centra  as  A.  calva  and  A.  scutata,  with  the 
number  of  fused  hypurals  also  essentially 
the  same  (Table  9).  Thus  A.  fragosa  and 
A.  uintaensis  are  meristically  distinct  from 
one  another  and  also  from  A.  scutata  and 
A.  calva,  suggesting  that  these  two  earlier 
forms  can  be  taxonomically  separated  on 
vertebral  meristic  characters. 

VERTEBRAL  COLUMN  OF  AMIA  CALVA 

The  existing  taxonomy  of  many  North 
American  fossil  amiids  is  based  primarily 
on  vertebral  characters.  Many  of  the  spe- 
cies of  "Protamia,"  and  the  genus  itself  as 
described  by  Leidy  (1873a)  from  the 
Bridger  Formation,  have  been  established 
solely  on  height/ width  proportions  and 
length  (thickness),  shape  of  the  neural 
and  aortal  facets,  and  various  foramina  of 
isolated  vertebrae.    Fossil  species  of  Amia 


from  the  Bridger  and  Cypress  Hills  forma- 
tions have  also  been  defined  on  character- 
states  of  isolated  vertebrae.  In  order  to 
analyze  this  usage,  variation  in  vertebral 
character-states  of  A.  calva  has  been  studied. 
The  axial  skeleton  of  Recent  Amia  calva 
is  relatively  well  known.  It  is  one  of  the  few 
modem  forms  that  have  diplospondylous 
vertebral  centra  posteriorly,  a  condition 
that,  according  to  Schaelfer  (1967),  func- 
tionally increases  the  flexibility  of  the  pos- 
terior part  of  the  body.  Shufeldt  (1885) 
was  one  of  the  first  to  describe  the  verte- 
brae of  Amia,  and  Hay's  ( 1895 )  well- 
known  work  on  the  vertebral  column  of 
Amia  provides  a  relatively  complete  and 
informative  description  of  the  axial  skele- 
ton, as  well  as  one  of  the  first  discussions 
of  intracolumnar  variation  of  the  centra. 
Hay  observed  some  gradual  changes  in 
centrum  proportions,  and  in  the  position  of 
the  neural  and  aortal  facets. 

Vertebral  Features 

Dorsal  and  ventral  facets,  basapophyses, 
foramina,  and  ridges  on  the  centra  have 
been  used  as  diagnostic  characters  in  the 
taxonomy  of  fossil  amiids.  There  are  three 
types  of  paired  facets  on  the  vertebrae: 
dorsal  neural  facets  for  the  neural  arches, 
ventral  aortal  facets  for  the  aortal  supports, 
and  haemal  facets  for  the  haemal  arches. 

Neural  facets.  The  neural  facets  are 
shallow  depressions  under  the  neural  arch 
bases,  which  in  life  are  filled  with  cartilage. 
Cartilage  is  present  between  the  centrum 
and  its  associated  neural  arch.  Some  speci- 
mens of  A.  calva  have  much  deeper  facets, 
with  a  small  ossified  ridge  built  up  on  the 
borders.  These  neural  facets  occur  in  pairs 
on  the  dorsal  surface  of  both  trunk  and 
caudal  vertebrae,  and  between  the  two 
facets  lies  a  groove  that  partially  receives 
the  spinal  cord. 

According  to  Hay  (1895:  7-9),  there  is  a 
marked  anteroposterior  change  in  the  posi- 
tion of  the  neural  facets.  He  contended 
that  at  the  anteriormost  end  of  the  vertebral 
column  the  neural  arch  bases  occur  be- 
tween two  vertebrae  and  rest  equally  on 


Fossil  Amiids  •  Boraske 


29 


VENTRAL 


dUJZ^ 


odZlL^ 


28 

35 
36 

37 
38 


4II> 


DORSAL 


Fig.    11.      Configuration    of    aortal    facets    (as)    and    neural    facets    (ns)    on    trunk    and    anterior    caudal    vertebrae    of 
Amio  calva  (339  mm  SL). 


both;  going  posteriorly  the.se  bases  shift 
gradually  backward.  He  also  observed  that 
there  is  a  change  in  the  spacing  of  the 
neural  iuches;  they  are  close  together  in  the 
anterior  trinik  region  and  more  widely 
spaced  posteriorly.  Hay  is  correct  in  regard 
to  the  change  in  spacing  of  the  neural 
arches,  but  he  is  not  altogether  correct  in 
his  description  of  the  change  in  position 
of  these  arches  in  relation  to  the  centra. 
An  examination  of  the  Wisconsin  A.  calva 
sample  showed  that,  after  the  first  few  an- 
teriormost  centra  and  corresponding  neural 
arches,  the  middle  of  the  neural  arches  is 
situated  at  the  juncture  between  the  centra. 
This  placement  continues  along  the  axial 
column  until  the  first  diplospondylous  ver- 


tebra occurs.  At  this  point,  the  next  five 
to  seven  neural  arches  are  found  aligned  to 
the  middle  of  each  of  the  corresponding 
centra,  after  which  the  arches  appear  to 
move  forward  slightly  and  correspond  ir- 
regularly to  the  vertebral  bodies. 

The  configuration  of  the  neural  facets 
themselves  varies  in  the  trunk  region  of  the 
vertebral  column  of  A.  calva.  The  neural 
arches  in  the  anterior  trunk  region  are 
thicker  and  wider  than  those  in  the  more 
posterior  trunk  region  which  have  become 
more  flattened  and  elongated.  The  shape 
of  the  neural  facets  reflects  this  trend  ( Fig. 
11).  After  the  first  two  ccMitra,  the  facets 
assume  an  hourglass  shape,  being  narrower 
in  the   middle   and  broader   at   each   end. 


30         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


This  can  be  related  to  the  fact  that  the 
neural  arches  are  situated  at  the  juncture  of 
two  centra  so  that  each  neural  facet  sup- 
ports the  anterior  and  posterior  halves  of 
two  different  neural  arches,  whose  bases 
are  narrow  at  the  extremity  and  thick  in 
the  center.  Although  the  neural  facets  in 
any  given  specimen  of  A.  calva  conform  to 
this  general  trend,  the  individual  configura- 
tion of  the  facets  varies  slightly.  Given  this 
variation  in  shape  of  the  neural  facets,  it  is 
useless  to  attempt  characterization  of  the 
vertebral  column  of  any  amiid  species  based 
on  configuration  of  neural  facets. 

Aortal  and  haemal  facets.  On  the  ven- 
tral side  of  the  trunk  vertebrae  are  two  thin 
cartilaginous  projections  that  are  located 
on  either  side  of  the  dorsal  aortal  supports. 
When  the  skeleton  is  dried,  these  projec- 
tions leave  marked  depressions,  which,  like 
the  neural  facets,  vary  gradually  from  the 
first  anterior  vertebra  to  the  last  few  trunk 
vertebrae;  at  this  point  the  aortal  facets 
coalesce  with  the  basapophyses  (Fig.  11). 
The  point  where  these  two  elements  are 
completely  merged  marks  the  termination 
of  the  trunk  centra,  and  the  next  centrum 
is  that  of  the  first  caudal  vertebra.  These 
structures,  which  were  derived  anteriorly 
from  the  basapophyses  and  aortal  facets, 
here  become  the  haemal  facets. 

The  first  pair  of  aortal  facets  is  very  small 
and  ovoid.  The  next  few  centra  bear  aortal 
facets  that,  as  Hay  (1895)  also  observed, 
are  circular.  The  following  aortal  facets 
become  successively  elongated,  until,  with 
the  tenth  or  twelfth  vertebra,  these  facets 
have  evolved  into  a  long  pair  of  slits,  usu- 
ally narrower  at  the  midpoint.  Posteriorly, 
these  slitlike  aortal  facets  remain  basically 
the  same  shape  until,  at  the  end  of  the 
dorsal  trunk  region,  they  merge  with  the 
basapophyses  to  form  haemal  facets.  Hay 
(1895:  54-57)  states  that  the  cartilaginous 
aortal  supports  penetrate  deeply  into  the 
centra  of  younger  individuals,  while  in 
older  specimens  they  rest  superficially  on 
the  centra.  The  aortal  facets  are  deeper 
and  more  distinct  than  the  neural  facets. 

Beginning  with  approximately  the  tenth 


or  twelfth  vertebra,  the  slit-shaped  aortal 
facets  are  vertically  situated  on  either  side 
of  an  indentation  that  contains  the  aorta 
(Fig.  11).  The  first  four  centra  have 
thicker  and  shorter  supports  with  relatively 
little  or  no  space  between  them.  The  aorta 
lies  ventrally  under  the  basioccipital, 
which  bears  aortal  supports  whose  facets 
are  of  the  same  shape  as  the  first  four  centra 
(Estes  and  Berberian,  1969,  fig.  2B  for 
A.  frafi^osa).  The  aortal  facets  of  the  first 
eight  vertebrae  are  different  from  all  other 
trunk  vertebrae,  whose  shape,  as  mentioned 
above,  is  basically  an  elongated  slit.  These 
aortal  supports  are  thus  helpful  in  dis- 
tinguishing the  first  eight  or  so  vertebrae 
from  the  remainder  of  the  trunk  centra  in 
disarticulated  specimens  (Fig.  11). 

Haemal  facets.  The  haemal  facets, 
which  contain  a  cartilaginous  layer  be- 
tween the  centrum  and  the  haemal  arches, 
are  nearly  rectangular-shaped  pairs  that  do 
not  vary  along  the  caudal  portion  of  the 
vertebral  column  until  the  first  fused  urals. 
The  furrow  or  indentation  that  lies  between 
the  aortal  facets  in  the  trunk  centra  con- 
tinues in  the  caudal  region  between  the 
paired  haemal  facets,  although  it  gradually 
decreases  in  width  and  depth.  Unlike  the 
neural  facets,  the  haemal  facets  are  out- 
lined by  an  ossified  border,  which  can  be 
helpful  in  distinguishing  dorsal  from  ventrid 
surfaces  in  disarticulated  caudal  vertebrae. 

Since  the  ossified  walls  are  tilted  20 
degrees  posteriorly  to  accommodate  the 
haemal  arches,  which  articulate  with  the 
cartilaginous  layer  diagonally  rather  than 
laterally,  those  borders  are  also  useful  in 
determining  the  anteroposterior  orientation 
of  the  centrum. 

Basapophyses.  Amia  trunk  centra  are 
distinguished  from  the  caudal  vertebrae  by 
their  having  prominent  paired  processes, 
which  ha\e  been  called  transverse  pro- 
cesses, parapophyses,  or  diapophyses.  I 
follow  the  terminology  of  Bolk  et  al.  ( 1936), 
wherein  they  designate  these  structures, 
which  are  the  processes  for  pleural  ribs,  as 
basapophyses  ( "basalstiimpfe" ) .  The  first 
centrum    often    lacks    these    basapophyses 


Fossil  Amiids  •  Boreske        31 


20 


44 


24 


48 


28 


56 


8 


32 


<k^ 


64 


36 


68 


40 

Fig.    12.      Shape  of  selected  trunk  and  caudal  vertebrae  in  Amia  calva  (339  mm  SL). 


78 


(Tables  10-12),  which  are  always  present 
on  the  succeeding  centra  and  progressively 
become  longer  until  appro.ximately  the 
twelfth  (Fig.  12).  The  basapophyses  are 
approximately  the  same  length  between 
the  twelfth  and  tlie  thirty-second  centra, 
from  which  point  they  begin  to  diminish 
gradually  in  length  until  the  last  trunk 
centrum,  where  they  coalesce  with  the 
aortal  facets.  The  lengths  of  the  basapo- 
physes were  not  individually  measured; 
this  data  would  be  of  little  practical  use  in 
a  comparison  of  Recent  and  fossil  material 
since  these  relatively  fragile  structures  are 
rarely  preserved  intact  in  fossils.  The  distal 
end  of  each  basapophysis  is  attached  to  a 
pleural    rib    by    means    of    cartilage.     The 


proximal  ends  of  the  basapophyses  are 
ankylosed  to  the  ventral  half  of  the  verte- 
bral body.  These  paired  processes  are  solid 
cyHnders  (hollow  at  the  tips)  that  are 
slightly  Battened  dorsoventrally.  Each  pair 
of  basapophyses  may  not  always  be  of  equal 
length  or  diameter,  but  they  are  extremely 
regular  in  position.  They  form  two  con- 
tinuous and  symmetrical  lines  that  gradu- 
ally come  closer  together  until  the  last 
trunk  centrum,  where  they  are  separated 
only  by  aortal  supports. 

An  important  aspect  of  the  basapophyses 
in  A.  calva  is  the  angle  between  each  indi- 
vidual pair  which  gradually  decreases  pos- 
teriorly. Since  the  angl(>  ])etween  the 
basapophyses  is  generally  still  available  in 


32        Bulletin  Museum,  of  Comparative  Zoology,  Vol.   146,  No.   1 


fossil  forms,  even  in  those  with  broken  basa- 
pophyses,  it  is  used  here  as  a  basis  of 
comparison  between  the  Recent  and  fossi! 
forms.  Since  the  angles  steadily  decrease 
posteriorly  along  the  vertebral  column 
(Figs.  12,  14),  they  are  also  useful  in 
orienting  disarticulated  centra  to  approxi- 
mate position  along  the  column.  Although 
there  is  individual  variation  in  these  angles 
(Tables  10-12),  they  are  nevertheless  con- 
sistent enough  to  help  in  determining  the 
general  position  in  the  column  of  any 
single  trunk  centrum.  The  range  of  angles 
extends  from  approximately  180  degrees 
anteriorly  to  45  degrees  posteriorly.  Since 
the  three  A.  calva  specimens  studied  were 
of  varying  sizes  ( 193  mm  SL,  382  mm  SL, 
and  423  mm  SL),  it  would  appear  that  there 
is  no  significant  change  in  the  angles  with 
increasing  size  or  age  of  the  fish  (Fig.  14). 
Although  this  transition  is  not  perfectly 
linear,  the  angles  are  always  decreasing 
posteriorly,  and  at  least  in  the  specimens  I 
measured,  there  was  never  an  instance  of 
an  angle's  measurement  being  greater  than 
that  of  the  preceding  centrum.  The  angle 
decrease  occurs  at  a  fairly  constant  rate 
until  approximately  the  thirtieth  trunk  ver- 
tebra, at  which  point  the  rate  of  decrease 
of  the  angles  is  much  accelerated  ( Fig.  14 ) . 
The  angle  of  the  basapophyses  is  thus  a 
reliable  parameter  in  identifying  the  gen- 
eral position  of  isolated  trunk  centra. 

Foramina,  bone  ridges,  and  first  centrum. 
The  trunk  centra  of  A.  calva  have  lateral 
foramina  that,  although  lacking  the  uni- 
formity of  the  neural  and  aortal  facets, 
occur  in  irregular,  distinct  paired  linear 
patterns.  The  foramina  of  the  tnmk  and 
caudal  vertebrae  transmit  numerous  small 
blood  vessels. 

On  the  lateral  surfaces  perpendicular  to 
the  anterior  and  posterior  articular  surfaces 
of  the  individual  centra  are  prominent  bone 
ridges.  These  bone  ridges  add  support  to 
the  arch  anlagen,  and  also  help  unify  the 
anlagen  into  a  sturdy,  functional  vertebral 
body  (Schaeffer,  1967).  Externally,  these 
bone  ridges  are  not  as  regular  as  they  are 
internally,    although   they   still   lie   antero- 


posteriorly  in  the  lateral  and  ventral  regions 
and  extend  vertically  along  the  basapo- 
physes. They  are  also  quite  prominent  in 
the  notochordal  furrow.  Such  bone  ridges 
are  not  a  unique  feature  of  A.  calva,  and  are 
common  in  teleosts. 

The  centra  in  A.  calva  are  amphicoelous. 
The  first  four  to  six  centra  differ  from  all 
corresponding  centra  by  having  the  anterior 
articular  surface  more  convex  than  concave. 
The  first  centrum  in  nearly  all  specimens 
observed  lacked  basapophyses,  and  should 
therefore  be  considered  a  minor  taxonomic 
character  since  first  centra  do  occasionally 
occur  with  very  small  basapophyses.    The 


Angle    of 
basapophyses 


Fig.    13.      Index  to  the   measurements   used,  superimposed 
upon   an   outline   drawing   of  an   Amia   calva   vertebra. 


Fossil  Amiids  •  Boreske        33 


^.  co/yo  verfebrol  lengths 


1(  24  n  <0  <l  Si  (4  72  10  II 

Verl.  no. 


A.calvo  bosopophyseol  angles 


12  It  21 

Vert.  ■•. 


---^ 


12  41  41 

Vtrt.  ■•. 


Vx 


y 


v^i 


•-v.' 


A.  uinfaensis 


NEIGIT 

mom 


I       i         ii        H        W 


—» — jr 

Vtrt.  M. 


Ti n w 


Fig.  14.  Intracolumnar  variation  in  the  angle  of  basapophyses,  length,  height,  and  width  of  vertebrae  in  Re- 
cent Ami'o  co/vo  (A  =  423  mm  SL;  B  ^  382  mm  SL;  C  =:  193  mm  SL).  Intracolumnar  variation  in  height  and  width 
of  trunk  and  caudal  vertebrae  in  A.  uin/oensi's.  Vertebral  column  model  based  on  first  six  centra  from  PL)  10101 
and  fifty-nine  centra  from  CM  25362;  missing  caudal  centra  have  been  interpolated  and  inferred  based  on  PL)  13865. 
The  first  anterior  centra  (PL)  10101)  were  larger  specimens  and  thus  the  anterior  region  of  the  trunk  vertebral  column 
model  is  "out-of-phase."  Vertical  lines  =  last  trunk  centrum. 


ovoid  shape  of  the  aortal  facets  is  a  constant 
feature  of  all  first  four  to  six  centra  ob- 
served. 

Vertebral  Dimensions 

A  superficial  but  often-used  character  for 
diagnosing  fossil  amiid  species  has  been  the 
shape  of  the  centrum.  Descriptions  for 
Amia  whiteavesiana,  A.  macrospondyla,  A. 
exilis,  A.  elegans,  A.  depressus,  A.  newher- 
rianus,  Protamia  symphysis,  P.  media,  P. 
gracilis,  P.  uintaensis,  P.  plicatus,  P.  cor- 
sonii,  and  P.  laevis  include  centrum  mea- 
surements for  height,  width,  and  length 
(thickness),  as  well  as  qualitative  descrip- 


tions of  the  fonii  and  proportions  of  the 
centrum.  Because  isolated  amiid  vertebrae 
have  often  been  the  only  anatomical  mate- 
rial found  in  the  fossil  record,  the  original 
diagnoses  were  obviovisly  limited  in  that  a 
great  deal  of  emphasis  was  placed  on  the 
vertebral  centrum.  In  considering  a  single 
centrum  shape  as  diagnostic  for  an  amiid 
species,  early  authors  implicitly  assumed 
the  vertebral  column  to  be  static,  with  no 
physical  change  or  variation  among  the 
centra  other  than  regional.  Many  new 
species  were  therefore  described  solely  on 
variation  in  shape  from  other  known  amiid 
types. 


34         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Table  10.     iNiTtAcoLUMNAR  variation  in  height,  width,  length,  and  angle  of  basapophyses 

OF  VERTEBRAE  IN  RECENT  Amiu  colva  ( 193  mm  SL) 


Vert. 
Cent. 


Height 
(mm) 


Width 
(mm ) 


Length 
(mm) 


Angle  of 
Basapophyses 
( Degrees ) 


Vert. 
Cent. 


Height 
(mm) 


Width 

(mm) 


Length 
(mm) 


1 

3.55 

4.35 

1.40 

44 

2.75 

2.75 

1.35 

2 

3.65 

4.20 

1.70 

176 

46 

2.60 

2.65 

1.15 

4 

3.70 

4.25 

1.65 

159 

48 

2.65 

2.65 

.90 

6 

3.40 

4.15 

1.70 

159 

50 

2.55 

2.60 

1.30 

8 

3.25 

3.90 

2.05 

153 

52 

2.50 

2.45 

1.35 

10 

3.15 

3.65 

2.00 

145 

54 

2.50 

2.50 

1.20 

12 

3.00 

3.65 

1.90 

140 

56 

2.45 

2.45 

1.15 

14 

3.05 

3.60 

2.00 

138 

58 

2.50 

2.45 

1.20 

16 

2.95 

3.55 

2.20 

134 

60 

2.30 

2.40 

1.15 

18 

3.00 

3.55 

2.30 

130 

62 

2.40 

2.30 

1.15 

20 

3.05 

3.40 

2.10 

122 

64 

2.30 

2.20 

1.15 

22 

2.90 

3.65 

2.25 

118 

66 

2.30 

2.05 

1.10 

24 

2.75 

3.25 

2.10 

103 

68 

2.25 

2.15 

1.00 

26 

3.00 

3.50 

2.15 

99 

70 

2.30 

2.15 

.80 

28 

3.05 

3.35 

2.20 

93 

72 

2.30 

2.10 

.85 

30 

3.00 

2.90 

2.25 

91 

74 

2.25 

2.05 

1.00 

32 

2.95 

3.05 

2.15 

90 

76 

2.05 

1.90 

.90 

34 

3.00 

3.00 

2.05 

74 

77 

1.87 

1.75 

.90 

36 

3.10 

3.08 

2.25 

63 

80 

1.65 

.80' 

37 

3.15 

3.00 

2.15 

44 

82 

1.50 

.68 

38 

3.25 

3.00 

2.10 

84 

1.25 

.53 

40 

2.80 

2.90 

2.25 

86 

.80 

.50 

42 

2.75 

2.90 

1.50 

"  Fused. 


Table  11.     Intracolumnar  variation  in  height,  vi^idth,  length,  and  angle  of  basapophyses 

OF  vertebrae  in  recent  Amia  calva  (382  mm  SL) 


Vert. 
Cent. 


Height 

(mm) 


Width 
(mm) 


Length 
(mm) 


Angle  of 

Basapophyses 

( Degrees ) 


Vert. 
Cent. 


Height 

(mm) 


Width 

(mm) 


Length 
(mm) 


1 

6.30 

7.21 

1.90 

180 

42 

5.50 

5.05 

3.50 

2 

6.25 

7.41 

2.45 

173 

44 

5.15 

4.65 

2.40 

4 

6.00 

7.50 

2.50 

161 

46 

5.00 

4.80 

2.35 

6 

5.95 

7.25 

2.85 

161 

48 

4.90 

4.60 

2.37 

8 

5.82 

7.10 

2.95 

159 

50 

4.70 

4.45 

2.50 

10 

5.56 

6.90 

3.05 

154 

52 

4.70 

4.45 

1.90 

12 

5.50 

6.65 

3.45 

143 

54 

4.65 

4.55 

2.00 

14 

5.50 

6.25 

3.45 

140 

56 

4.70 

4.45 

2.10 

16 

5.50 

6.25 

3.45 

133 

58 

4.80 

4.10 

1.90' 

18 

5.35 

6.30 

3.50 

125 

60 

4.40 

4.45 

2.00 

20 

5.25 

6.25 

3.55 

120 

62 

4.80 

4.45 

2.12' 

22 

5.30 

6.25 

3.55 

116 

64 

4.50 

4.10 

2.12' 

24 

5.30 

6.25 

3.65 

113 

66 

4.40 

3.85 

2.00 

26 

5.20 

6.20 

3.85 

110 

68 

4.00 

3.75 

2.00 

28 

5.35 

6.15 

3.70 

110 

70 

4.05 

3.70 

1.87 

30 

5.25 

6.00 

3.60 

105 

72 

3.75 

3.50 

1.75 

32 

5.35 

5.80 

3.85 

100 

74 

3.50 

3.35 

1.50 

34 

5.45 

5.50 

3.90 

92 

76 

3.20 

3.15 

1.47 

36 

5.50 

5.15 

3.50 

78 

78 

2.65 

1.65 

38 

5.75 

5.10 

3.55 

66 

80 

2.00 

1.40 

39 

5.70 

5.07 

3.60 

46 

82 

1.95 

1.20 

40 

5.65 

5.05 

3.65 

84 

1.80 

1.12 

•  Fused. 


Fossil  Amiids  •  Boreske        35 


Table  12.     Intracolumnar  variation  in  height,  width,  length,  and  angle  of  basapophyses 

OF  vertebrae  in  recent  Amia  calva  (423  mm  SL) 


Vert. 
Cent. 


Height 
(mm) 


Width 

(mm) 


Length 
(mm) 


Angle  of 

Basapophyses 

( Degrees ) 


Vert. 
Cent. 


Height 
(mm ) 


Width 
(mm) 


Length 
(mm) 


1 

8.25 

9.25 

2.80 

44 

6.45 

6.40 

3.45 

2 

8.00 

9.15 

3.35 

172 

46 

6.40 

6.21 

3.05 

4 

7.80 

9.05 

3.40 

156 

48 

6.20 

6.00 

3.05 

6 

7.55 

9.20 

3.45 

150 

50 

6.10 

5.85 

3.85 

8 

7.45 

8.80 

3.45 

142 

52 

6.25 

5.85 

3.80 

10 

7.40 

8.80 

3.75 

140 

54 

6.10 

5.85 

2.95 

12 

7.25 

8.75 

3.85 

132 

56 

5.95 

5.90 

2.70 

14 

7.25 

8.75 

4.00 

126 

58 

5.80 

5.85 

2.65 

16 

7.35 

8.70 

4.05 

119 

60 

5.80 

5.65 

2.60 

18 

7.15 

8.50 

4.15 

117 

62 

5.65 

5.70 

2.80 

20 

7.15 

8.50 

4.10 

114 

64 

5.60 

5.30 

2.70 

22 

7.15 

8.40 

4.00 

112 

66 

5.55 

5.20 

2.45 

24 

7.35 

8.35 

4.37 

103 

68 

5.50 

5.05 

2.50 

26 

7.45 

8.30 

4.75 

103 

70 

5.30 

4.95 

2.35 

28 

7.30 

8.20 

4.75 

95 

72 

5.30 

4.90 

2.20 

30 

7.47 

8.10 

4.70 

90 

74 

5.20 

4.80 

2.10 

32 

7.25 

7.95 

4.95 

90 

76 

5.05 

4.65 

1.85 

34 

7.35 

7.45 

5.00 

67 

78 

5.00 

4.50 

2.10* 

36 

7.35 

6.90 

5.00 

52 

80 

4.75 

4.45 

2.25 

37 

7.50 

6.70 

4.95 

46 

82 

4.15 

1.55 

38 

7.65 

6.50 

4.90 

84 

3.25 

1.50 

40 

7.60 

6.50 

4.50 

86 

2.50 

1.50 

42 

7.40 

6.20 

4.35 

88 

2.00 

1.45 

<•  Fused. 


One  of  the  detailed  studies  on  intra- 
columnar vertebral  variation  is  Hoffstetter 
and  Case's  ( 1969 )  work  on  the  vertebral 
column  of  snakes.  Measuring  individual 
centra  in  sequence,  they  plotted  this  varia- 
tion; similar  graphs  are  used  here  (Fig. 
14 ) .  Three  specimens  of  A.  calva  ( 193 
mm  SL,  382  mm  SL,  and  423  mm  SL )  were 
dissected  and  the  individual  vertebral 
dimensions  measured  to  determine  verte- 
bral variation. 

Length.  Centrum  length  (thickness) 
was  measured  anteroposteriorly  at  the  mid- 
line, above  the  basapophyses  (Fig.  13). 
It  was  necessary  to  establish  such  a  control 
for  this  measurement  because  of  the  varia- 
tion in  thickness  within  each  centrum.  In 
the  trunk  region,  the  centra  are  thickest 
ventrally  at  the  neural  and  dorsal  facets. 
The  caudal  vertebrae  follow  a  similar  pat- 
tern, being  slightly  thicker  ventrally  and 
dorsally,  and  thinner  laterally.  Every 
second  centrum  was  measured  for  length 
(Tables  10-12).    There  is  a  distinct  intra- 


columnar variation  for  this  measurement, 
although  the  difference  in  length  between 
consecutive  vertebrae  is  usually  small. 
There  is  also  a  general,  if  somewhat  ir- 
regular, pattern  in  centrum  length  in  A. 
calva  (Fig.  14).  The  first  two  or  three  ver- 
tebrae of  each  A.  calva  specimen  are  rela- 
tively thin.  These  are  followed  by  centra 
that  gradually  increase  in  length  until  ap- 
proximately the  last  trunk  centrum  at  the 
midbody.  At  this  point  there  is  a  general 
trend  again  towards  thinner  vertebrae,  al- 
though this  pattern  is  erratic,  particularly 
between  the  fiftieth  and  the  fifty-fourth 
centra,  where  the  thickness  is  suddenly  in- 
creased and  then  decreases.  This  sudden 
change  here  in  vertebral  thickness  occurs 
directly  above  the  midline  of  the  anal  fin. 
The  shortest  vertebrae  are  the  fused  urals. 
Height.  Centrum  height  was  taken  dor- 
soventrally  at  the  midline,  between  the 
aortal  and  neural  facets  (Fig.  13).  Every 
second  centrum  was  measured  up  to  the 
fused  urals,  in  which  an  accurate  measure- 


36 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Table  13.     Intracolumnar  variation  in  height,  width,  length,  and  angle  of  basapophyses 

OF  VERTEBRAE  IN  Atnla  uintaensis  CM  25362 


Vert. 
Cent. 


Height 
(mm) 


Width 
(mm ) 


Length 

( mm ) 


Angle  of 

Basapophyses 

( Degrees ) 


Vert. 
Cent. 


Height 
(mm) 


Width 

( mm ) 


Length 
( mm ) 


7 

24.5 

32.8 

9.8 

179 

37 

19.5 

19.5 

7.5 

8 

25.0 

32.5 

9.8 

178 

38 

19.8 

18.8 

7.0 

9 

26.8 

32.0 

9.5 

177 

39 

19.8 

18.5 

6.9 

10 

26.5 

32.1 

9.5 

176 

40 

20.0 

17.5 

6.5 

11 

25.5 

32.8 

9.0 

174 

41 

— 

6.5 

12 

26.0 

31.0 

9.1 

172 

42 

19.5 

18.0 

6.5 

13 

25.4 

31.1 

9.5 

164 

43 

18.5 

17.5 

6.0 

14 

27.5 

31.0 

9.5 

160 

44 

18.5 

17.5 

6.2 

15 

25.5 

30.0 

10.0 

156.5 

45 

17.5 

17.5 

6.0 

16 

26.0 

30.5 

9.8 

154 

46 

17.2 

16.0 

6.0 

17 

26.5 

31.5 

9.5 

153 

47 

17.0 

16.0 

6.0 

18 

27.0 

30.5 

9.8 

149 

48 

16.5 

16.0 

5.5 

19 

25.1 

29.5 

9.5 

143 

49 

16.5 

— 

6.0 

20 

24.5 

29.5 

9.5 

132 

50 

16.2 

15.5 

5.5 

21 

24.0 

30.0 

9.5 

122 

51 

16.0 

15.0 

5.8 

22 

24.5 

28.2 

9.0 

117 

52 

16.0 

15.5 

5.0 

23 

24.0 

29.0 

9.5 

110 

53 

15.5 

14.0 

5.5 

24 

24.0 

28.5 

9.0 

102 

54 

15.2 

14.5 

5.2 

25 

24.0 

26.5 

8.5 

102 

55 

15.5 

13.2 

6.0 

26 

24.5 

27.0 

10.0 

97 

56 

15.0 

12.2 

5.5 

27 

23.5 

25.0 

11.0 

90 

57 

14.5 

12.0 

6.5 

28 

23.5 

25.0 

11.0 

83 

58 

14.5 

12.5 

5.5 

29 

23.2 

25.0 

12.0 

80 

59 

14.0 

11.5 

5.2 

30 

23.0 

22.5 

11.0 

62 

60 

14.0 

5.0 

31 

25.0 

22.5 

11.0 

46 

61 

13.5 

11.5 

5.0 

32 

— 

11.0 

62 

13.5 

10.5 

4.5 

33 

— 

10.0 

63 

4.5 

34 

23.0 

19.5 

9.5 

64 

12.0 

9.0 

4.5 

35 

22.5 

9.5 

65 

8.5 

7.0 

4.2 

36 

21.0 

20.0 

7.5 

ment  would  be  obscured  by  the  fusion  of 
the  hypurals.  This  series  of  measurements 
shows  a  basic  pattern  that  is  similar  for 
each  of  the  individuals  studied,  although 
there  is  less  intracolumnar  variation  in  the 
height  than  in  the  length  measurements 
(Tables  10-12).  The  greatest  height  gen- 
erally occurs  at  the  anteriormost  region  of 
the  column,  then  decreases  slightly  until 
the  midtrunk  region  (Fig.  14).  At  this 
point  the  height  gradually  increases  until 
it  peaks  at  the  end  of  the  trunk  region  and 
the  beginning  of  the  caudal  section,  after 
which  it  decreases  again  toward  the  caudal 
region  (Fig.  14). 

Width.  The  width  measurements  were 
taken  perpendicular  to  the  height  measure- 
ments, at  the  widest  section  of  the  centrum 
(Fig.  13).    This  dimension  has  a  greater 


linear  slope  than  the  length  and  height 
dimensions  ( Fig.  14 ) ,  which  follow  a  more 
bell-shaped  curve.  There  is  a  greater  varia- 
tion within  the  vertebral  column  for  width 
dimensions  (Fig.  14),  as  comparison  of  the 
height  and  width  slopes  reveals.  The  great- 
est intracolumnar  width  is  always  at  the 
anteriormost  portion  of  the  trunk  region, 
after  which  this  dimension  gradually  de- 
creases. There  appear  to  be  two  areas 
where  the  rate  of  decrease  is  greater,  these 
being  at  the  terminus  of  the  trunk  region 
and  at  the  first  fused  ural. 

Height/ width  ratio.  The  centrum  height/ 
width  ratio  has  been  a  commonly  used  diag- 
nostic character  in  amiid  taxonomy.  Hay 
(1895:  7)  correctly  noted  that  the  trunk 
vertebrae  are  somewhat  broader  than  high 
( Fig.  14 ) ,  and  at  the  terminus  of  the  trunk 


Fossil  Amiids  •  Boreske 


37 


region  the  centra  are  nearly  circular.  The 
proportions  tend  to  be  reversed  in  tlie 
eandal  region,  however,  with  the  height 
generally  exceeding  the  width,  although  to 
a  lesser  degree  than  the  proportional  dif- 
ference in  the  trunk  region.  The  basic  trend 
in  shape  through  the  vertebral  column  is 
tluis  a  marked  horizontally  elliptical  cen- 
trum approaching  a  progressively  circular 
one,  which  then  again  becomes  slightly 
vertically  elliptical.  Thus  there  is  quite  a 
variation  in  the  centrum  shape  throughout 
the  axiiil  column,  so  that  no  one  shape  or 
ratio  of  dimensions  could  reasonably  be 
considered  diagnostic  for  all  the  centra  of 
the  vertebral  column. 

VALID  NORTH  AMERICAN  FOSSIL 
GENERA  AND  SPECIES 

Amia  fragosa  (Jordan,  1927) 

Kitidleia  fragosa  Jordan,  1927:   145. 
SUjlomylcodon  lacus  Ru.ssell,  1928a:   103. 
Paramiatus  giirleyi  Ronier  and  Fryxell,  1928:  519. 

Holotype.  NMC  8533e,  anterior  portion 
of  right  dentary. 

Paratypes.  NMC  8534a-d,  f-n.  (a),  left 
operculum  fragment;  ( b ) ,  cranial  fragment; 
(c),  anterior  portion  of  left  dentary;  (d), 
anterior  portion  of  right  dentary;  (f-g), 
coronoid  with  two  styliform  teeth  preserved; 
(h-i),  vomer  without  teeth  preserved;  (j), 
styHform  tooth  fragment;  (k-1),  posterior 
portion  of  right  dentary;  ( m ) ,  left  maxilla; 
(n),  anterior  portion  of  right  dentary. 

Ttjpe  locality  and  horizon.  Rumsey,  Al- 
berta. East  half  of  section  31,  T  34  S,  R  21 
W,  Rumsey  Quadrangle,  Alberta;  Edmon- 
ton Formation. 

Age  range.  Campanian  (Late  Creta- 
ceous) to  Bridgerian  (Middle  Eocene). 

Hypodiii^m.  Cretaceous.  Oldman  For- 
mation, Alberta:  AMNH  5934,  palatal  frag- 
ments with  styliform  teeth;  AMNH  5935, 
operculum  and  dentary.  "Mesaverde"  For- 
mation, Wyoming:  AMNH  5932,  dentary 
and  numerous  coronoid  teeth;  AMNH  5933, 
vertebrae.  Judith  River  Formation,  Mon- 
tana: AMNH  10109,  left  vomer  bearing 
styHform  teeth;  AMNH  10110,  dentary  frag- 


ments, vertebrae,  and  skull  elements.  Ed- 
monton Formation,  Alberta:  ROM  .3064, 
coronoid  teeth;  ROM  3065,  dentaries,  verte- 
brae, and  cranial  fragments;  UA  5398-5507, 
articulated  and  disarticulated  specimens 
(see  O'Brien,  1969  for  identifications). 
Lance  Formation,  Wyoming:  AMNH  9316, 
pterotic;  AMNH  9315,  operculum;  CM 
25363,  dentaries;  PU  17013,  dentaries; 
UCMP  54013-54015,  54017,  54019,  54021- 
54030,  54035-54038,  54040-54056,  54059- 
54069,  54070-54120,  54141-54167,  54174- 
54180,  54188-54198,  54260,  54262,  dis- 
articulated elements  (see  Estes,  1964  for 
identifications).  Hell  Creek  Formation, 
Montana:  PU  17016,  17048,  dentaries;  PU 
17014,  coronoid  teeth;  PU  20554,  dentary 
and  vertebrae;  MCZ  9286-9293,  9390-9432, 
9559,  disarticulated  elements  ( see  Estes  and 
Berberian,  1969  for  identifications ) . 

Paleocene.  Fort  Union  Formation,  Wy- 
oming: PU  17115,  coronoid  teeth;  PU  17126, 
coronoid  teeth  and  vertebrae;  PU  17117, 
dentary  and  maxilla;  PU  21525,  portion  of 
cranial  roof  with  associated  dentaries;  PU 
20523,  dentary  and  coronoid  teeth;  PU 
21174,  vertebrae.  Paskapoo  Formation,  Al- 
berta: UA  131,  dentary,  numerous  tooth 
plates,  and  vertebrae.  Tongue  River  For- 
mation, Montana:  PU  20577,  vertebrae, 
premaxillary  fragment,  and  coronoid  teeth; 
PU  20578,  basioccipital  and  vertebrae;  PU 
17068,  vertebra  and  dentary  fragment. 
Melville  Formation,  Montana:  AMNH  2635, 
cranial  elements  and  associated  dentaries. 
Tullock  Formation,  Montana:  PU  17069, 
vomers. 

Eocene.  Will  wood  Formation,  Wyo- 
ming: MCZ  9264,  nearly  complete  skull; 
PU  18780,  tooth  plate;  PU  21175,  dentary 
fragment  and  coronoid  teeth;  PU  16756, 
dentary  and  cranial  fragments;  PU  17649, 
anterior  portion  of  skull;  PU  21173,  skull 
fragments  and  vertebrae;  PU  13261-13262, 
cranial  fragments  and  coronoid  tc>eth. 
Golden  Valley  Formation,  North  Dakota: 
PU  18567,  coronoid  teeth  and  vertebrae. 
Wasatch  Formation,  Wyoming:  PU  13260, 
tooth  plates;  PU  13259,  cranial  fragments 
and    dentaries.     Bridger    Formation    Wyo- 


B 


Fig.  15.  A,  Amia  calva.  Recent,  Wisconsin;  above,  lateral,  and  below,  dorsal  views  of  skull.  B,  Amia  scutafa.  Early 
and  Middle  Oligocene;  above,  lateral,  and  below,  dorsal  views  of  skull  (sensory  canal  system  and  pit-lines  are  not 
known  since  skull  elements  are  in  articulation).  C,  Amio  fragosa,  Late  Cretaceous  to  Middle  Eocene;  above,  lateral, 
and  below,  dorsal  views  of  skull  (sensory  canal  system  and  pit-lines  after  Estes,  1964).  D,  Amia  uinfaensis,  Paleocene 
to  Early  Oligocene;  above,  lateral,  and  below,  dorsal  views  of  skull  (sensory  canal  system  is  only  known  in  the 
mandible,  operculum,  nasal,  lacrimal,  antorbital,  extrascapular,  and  suprascapular,  all  of  which  conform  with  those 
of  A.  calva). 

Abbreviations:  a,  angular;  ao,  antorbital;  br,  branchiostegal  rays;  d,  dentary;  ds,  dermosphenotic;  es,  extrascapular; 
fr,  frontal;  io,  interoperculum;  io-  io''  io'*  io'^',  infraorbital  series  (suborbitals  &  postorbifals);  la,  lacrimal;  m, 
maxilla;  n,  nasal;  op,  operculum;  p,  preoperculum;  pa,  parietal;  pt,  pterotic;  r,  rostral  (ethmoid);  s,  suprascapular;  so, 
surangular;  sm,  supramaxilla;  so,  suboperculum.  Dotted  lines  indicate  the  sensory  canal  system;  dashed  lines  indi- 
cate pit-lines. 


38 


Fossil  Amiids  •  Boreske 


39 


ining:  YPM  6245,  vomer  and  cranial  frag- 
mcMiVs;  YPM  6246,  vertebrae;  YPM  6247, 
dentary;  YPM  6248,  vertebra  and  cranial 
fragments;  YPM  6254,  verte]:)rae,  basioccip- 
ital,  vomer;  YPM  6261,  left  opercnlnm; 
ANSP  5630,  vertebra.  Green  RivcT  Forma- 
tion, Wyoming:  MCZ  5341,  FMNH  2201, 
complete  specimens. 

Known  distribution.  North  Dakota, 
Wyoming,  Montana,  and  Alberta. 

Revised  diapwsis.  Vertebral  colimm 
with  significantly  fewer  total  ctMitra  (65 
mean)  than  the  other  species,  with  approxi- 
mately twelve  fewer  trnnk  vertebrae  (25 
mean)  and  eight  fewer  monospondylous 
caudal  centra  ( 17  mean ) .  Distance  be- 
tween anal  fin  insertion  and  the  end  of  the 
vertebral  column  relatively  short,  with 
dorsal  fin  terminating  close  to  caudal  fin. 
Caudal  lepidotrichia  19-20  rather  than  23- 
27.  Ascending  processes  of  parasphenoid 
perpendicular  to  the  main  anteroposterior 
parasphenoid  axis;  more  posterior  place- 
ment of  parasphenoid  tooth-patch.  Pari- 
etals  squared  in  outline.  Marginal  teeth 
simple  pointed  cones,  palatal  teeth  usually 
stout  styliform  crushers.  Supraorbital  sen- 
sory canal  not  entering  parietal.  Excava- 
tion of  orbital  notch  in  frontal  relatively 
larger.  Dentary  with  additional  horizontal 
shelf  of  coronoid  articulation  surface  adja- 
cent to  lingual  border  of  alveolar  ridge; 
coronoid  articulation  surface  extensive, 
overlapping  ventral  half  of  ramus;  dentary 
with  pronoimced  arch  rather  than  gradual 
curve  in  ventral  outline.  Greatest  known 
standard-length  510  mm. 

Introduction 

Jordan  (1927)  described  Kindleia  fra^osa 
as  a  new  genus  of  cichlid  fish  from  the  Late 
Cretaceous  Edmonton  Formation  of  Al- 
berta. This  tentative  placement  of  Kindleia 
within  the  Cichlidae  was  largely  the  result 
of  his  misinteipreting  the  splenial  tooth 
plates  for  fused  lower  pharyngeal  bones 
(Estes,  1964).  One  month  later,  Russell 
( 1928a)  independently  published  a  descrip- 
tion of  Stylomyleodon  lacus,  a  new  fossil 
amiid  from  the  Late  Paleocene   Paskapoo 


Formation  of  Alberta,  and  referred  other 
specimens  from  the  Edmonton  Formation 
of  Alberta  to  the  same  species.  His  descrip- 
tion also  included  a  dentary  and  palatal 
teeth  modified  tor  crushing.  His  relegation 
of  the  genus  to  the  Amiidae  was  based  on 
a  correct  inteipretation  of  the  "splenial" 
(=  coronoid)  tooth  plates  (Estes,  1964). 
He  suggested  a  relationship  of  Stylonujle- 
odon  to  Platacodon  nanus  ( at  that  time 
erroneously  considered  an  amiid;  see  Estes, 
1964)  with  the  essential  difference  being 
hemispherical  rather  than  Hattened  tooth 
crowns. 

Jordan  later  ( 1928 )  noted  the  similarity 
of  the  two  genera  Kindleia  and  Stylomyle- 
odon and  asserted  the  prior  claim  of  his 
name  Kindleia.  Although  he  made  no  com- 
ment on  Russell's  attributing  Stylomyleodon 
to  the  Amiidae,  he  rejected  Russell's  com- 
parison of  that  genus  with  Platacodon  on 
the  basis  of  Marsh's  earlier  conviction  that 
the  latter  was  mammalian.  In  reply  to 
Jordan,  Russell  ( 1928b )  defended  the  valid- 
ity of  his  genus  on  the  supposition  that  its 
dentary  was  distinct  from  that  of  Kindleia, 
although  he  did  agree  on  the  similarity  of 
teeth  and  jaw  fragments  of  the  two  genera. 
Russell  (1929)  further  attempted  to  vali- 
date Stylomijleodon  as  a  genus  by  com- 
paring his  type  with  new  specimens 
collected  by  Princeton  University.  This  new 
material  confirmed  his  association  of  the 
maxilla-dentary  and  palatine-coronoid  den- 
titions, and  also  substantiated  his  interpreta- 
tion of  Stylomyleodon  as  an  amiid  in  which 
the  coronoid  teeth  were  specialized  for 
crushing.  He  also  admitted  that  there  was 
insufficient  Platacodon  material  to  deter- 
mine any  conclusive  similariti(\s  with  Stylo- 
myleodon, but,  r(>f(Mring  to  Hatcher's  (1900, 
1901 )  work,  did  insist  that  Platacodon  was 
a  fish.  Simpson  (1937)  reported  finding 
additional  specimens  of  Stylomyleodon  Rus- 
sell in  the  Fort  Union  Formation  at  Crazy 
Mountain  Field  sites  of  Montana. 

Estes  ( 1964 ) ,  from  his  studies  of  amiid 
material  from  the  Lance  Formation  of  Wyo- 
ming, observed  that  whereas  the  type 
dentary  referred  by  Russell  to  Stylomyleo- 


40 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


don  was  the  pcsterior  portion  of  an  amiid 
dentary,  Jordon's  type  was  the  anterior 
portion.  From  this  fact  he  confirmed  the 
synonymy  of  Stylomtjleodon  with  the  genus 
Kindleia,  at  that  time  beUeving  that  it  was 
generically  separated  from  Amia.  Janot 
( 1967 )  agreed  with  Estes  on  the  synonymy 
of  Stylomijleodon  with  Kindleia,  but  did  not 
find  sufficient  cause  to  distinguish  Kindleia 
generically  from  A7?j/fl.  Russell  (1967)  con- 
tinued to  leave  the  nomenclatural  problem 
of  Stylomyleodon-Kindleia  unsettled.  Estes 
and  Berberian  ( 1969)  studied  material  from 
the  Late  Cretaceous  Hell  Creek  Formation 
of  Montana  and  confirmed  Janot's  proposi- 
tion that  Kindleia  is  a  synonym  of  Amia. 
They  also  suggested  the  possibility  of 
synonymy  of  A.  fra<i,osa  with  A.  keJireri 
(Middle  Eocene,  Germany),  A.  russelli 
(Late  Paleocene,  France),  A.  munieri 
(Early  Oligocene,  France),  and  Paromiatus 
gurleyi  (Early  Eocene,  Wyoming),  but 
postponed  formal  synonymy  of  A.  jrag,osa 
with  the  prior  name  A.  kehreri  (Andreae, 
1892),  pending  more  detailed  study  of 
Early  and  Middle  Cenozoic  specimens 
from  Europe.  Estes  and  Berberian  (1969: 
10)  concluded  that  the  minor  variations 
that  separated  A.  fra<^osa  and  its  related 
forms  in  Europe  from  A.  calva  are  "super- 
ficial and  essentially  primitive,"  and  indi- 
cated that  the  group  might  be  close  to  the 
ancestry  of  the  Recent  species  A.  calva. 

A  nearly  complete  skull  from  the  Eocene 
Willwood  Formation  of  Wyoming  (Fig. 
16),  two  axial  skeletons  from  the  Eocene 
Green  River  Formation  of  Wyoming  ( Plate 
1 ) ,  and  a  sample  of  disarticulated  elements 
from  the  Late  Cretaceous,  Paleocene,  and 
Early  Eocene  have  yielded  more  informa- 
tion on  the  osteology  of  Amia  fragosa.  Estes 
(1964),  O'Brien  (1969),  and  Estes  and 
Berberian  (1969)  have  studied  this  species 
in  detail,  and  I  therefore  discuss  these  speci- 
mens only  as  they  modify  conclusions 
reached  by  those  studies. 

Fossil  Record 

In  addition  to  the  stratigraphic  list  given 
by  Estes  and  Berberian  (1969:  14,  table  1) 


of  major  freshwater  deposits  carrying  A. 
fragosa,  three  new  localities  are  recorded 
here:  the  Late  Paleocene  Silver  Coulee 
local  fauna  of  the  Fort  Union  Formation, 
Wyoming,  and  the  Early  Eocene  Willwood 
and  Wind  River  formations,  Wyoming.  The 
luajor  deposits  in  which  remains  of  A. 
fragosa  have  been  found  are  summarized  in 
Table  18.  Estes  and  Berberian  (1969:  10) 
state  that  the  stratigraphic  range  of  A. 
fragosa  extends  from  the  Late  Cretaceous 
through  at  least  Middle  Eocene  time  in 
North  America.  The  earliest  deposit  in 
which  remains  of  A.  fragosa  have  been 
found  is  the  Late  Cretaceous  (Campanian) 
Oldman  Formation  of  Alberta,  and  the 
latest  deposit  is  the  Middle  Eocene  (Brid- 
gerian)  Bridger  Formation  of  Wyoming. 
Cavender  (1968:  128),  however,  de- 
scribes Amia  scales  from  the  Late  Eocene 
( Duchesnean )  Clarno  Formation  of  Ore- 
gon. Although  these  small  scales  (approxi- 
mately 2  mm  in  length)  are  not  as  robust 
as  those  of  A.  fragosa,  they  are  more  ossified 
than  A.  scutata  and  A.  calva  scales.  These 
scales,  along  with  the  scales  from  the  Horse- 
fly River  Beds  of  British  Columbia  (UMMP 
collections)  cannot  at  present  be  identified 
as  to  species.  They  are  best  referable  to 
Amia  sp.,  since  no  identifiable  A.  fragosa 
elements  have  been  found  later  than 
Bridgerian  and  no  specific  character-states 
for  scales  of  A7nia  have  yet  been  deter- 
mined. 

Description 

Neurocranium.  Estes  (1964:  29)  stated 
that  the  greater  length  of  the  basioccipital 
and  the  presence  of  a  second  pair  of  aortal 
supports  in  Amia  calva  indicated  that  the 
basioccipital  posterior  to  the  spinal  (inter- 
vertebral) arterial  foramina  included  only 
one  fused  vertebra  in  Amia  fragosa  instead 
of  the  two  found  in  A.  calva.  O'Brien 
(1969:  42)  observed  a  similar  condition  in 
two  complete  A.  fragosa  specimens  from 
the  Edmonton  Formation  of  Alberta.  Estes 
and  Berberian  (1969:  2-3)  found  nine 
basioccipitals  with  one  fused  vertebra  and 
eleven  with  two  fused  vertebrae  from  the 


V 


Fossil  Amiids  •  Boreske        41 


Fig.    16.      Amia  fragosa  MCZ  9264,  Early  Eocene,  Willwood  Formation,  Wyoming;  A,  dorsal;  B,  ventrol. 


Hell  Creek  Formation  of  Montana  and  in- 
terpreted this  as  a  variation  in  A.  frci'^osa 
not  observed  in  the  Lance  sample.  Janot 
( 1967 )  noted  that  basioccipitals  of  the 
European  Late  Paleocene  Amia  sp.  also 
showed  this  variation.   Estes  and  Berberian 


(1969:  2)  suggested  that  a  weak  tendency 
for  fusion  of  vertebrae  could  be  correlated 
with  increasing  size,  and  that  such  varia- 
tion might  possibly  e.xist  in  the  Recent 
species  as  well.  Fifty  Recent  A.  calvo  skele- 
tons examined,  with  a  size  range  of  100-480 


42         Bullctiti  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


mm  SL,  have  the  first  two  vertebrae  fused 
to  the  basioceipital.  Three  articulated  and 
twenty-two  disarticulated  Eocene  and 
Oligocene  Amia  uintaensis  Ijasioccipitals 
all  have  two  vertebrae  fused  to  the  basioc- 
eipital. Unfortunately,  in  specimens  of  A. 
fni'^osa  (MCZ  9264,  PU  13261)  having  a 
visible  parasphenoid,  the  basioceipital  re- 
gions are  poorly  preserved.  There  is  a  pos- 
sibility that  the  Lance  sample  by  chance 
contained  only  specimens  with  one  fused 
vertebra  since  there  are  only  six  specimens 
known.  Until  more  specimens  of  A.  jra<iosa 
and  A.  uintaensis  with  intact  basioccipitals 
become  available,  it  is  difficult  to  discuss 
this  point  further. 


Fig.  17.  Comparison  of  parasphenoids  of  Amio  spp.: 
c,  Amia  calva.  Recent,  after  Janot,  1967  (c''  :=  dorsal,  c^' 
=  ventral);  f,  A.  fragosa  (ventral);  u,  A.  uinfaensis 
(ventral). 


The  length  of  the  A.  fra<j,osa  parasphe- 
noid posterior  to  the  ascending  processes  is 
10  percent  shorter  and  slightly  wider  than 
in  A.  calva,  with  the  ascending  processes 
more  posterior  than  in  the  Recent  species 
(Fig.  17).  The  proportion  of  the  length 
posterior  to  the  processes  to  the  length 
anterior  to  these  processes  (0.780)  is  not 
as  small  as  in  A.  uintaensis  (0.704)  or  as 
great  as  in  A.  calva  (0.900),  and,  on  the 
basis  of  this  small  sample,  A  fragosa  is  inter- 
mediate among  the  three  species  for  this 
character.  The  region  posterior  to  the 
processes  also  appears  more  convex  than  in 
A.  calva,  but  not  as  convex  as  in  A.  uijitaen- 
sis.  The  ascending  processes  are  almost 
perpendicular  to  the  main  anteroposterior 
axis  of  the  parasphenoid.  Those  of  A. 
fragosa  form  an  approximately  85-degree 
angle  with  the  parasphenoid  axis,  while  the 
ascending  processes  of  both  A.  calva  and 
A.  uintaensis  form  approximately  70-75- 
degree  angles.  The  mid-ventral  surface  of 
the  parasphenoid  bears  small,  sharp,  con- 
ical teeth.  This  tooth-bearing  surface  of  A. 
fragosa  terminates  anteriorly  toward  the 
middle  of  the  ascending  processes,  whereas 
in  A.  calva  this  region  narrows  to  a  point 
and  extends  to  the  posterior  end  of  the 
vomers  (Fig.  17).  In  A.  uintaensis,  this 
region  also  extends  to  the  vomers,  but 
covers  a  wider  surface  area  in  the  anterior 
region  than  in  A.  calva.  Nearly  all  the 
tooth-bearing  surface  of  A.  frasj^osa  lies  in 
the  posterior  half  of  the  parasphenoid, 
while  in  A.  calva  this  surface  is  centered 
between  the  posterior  and  anterior  areas; 
in  A.  uintaensis  two-thirds  of  this  surface 
lie  in  the  anterior  region  of  the  para- 
sphenoid. The  entire  tooth-bearing  surface 
of  A.  fragosa  is  wider  than  that  of  A.  calva, 
since  the  anterior  half  of  the  tooth-bearing 
surface  tapers  anteriorly  in  A.  calva,  while 
in  A.  fragosa  the  anterior  portion  maintains 
a  more  constant  width.  The  basic  outline  of 
the  tooth-bearing  surface  in  A.  fragosa  is 
subrectangular;  that  of  A.  calva  is  more 
tear-drop  shaped,  with  the  anterior  apex 
widened  and  extended  to  the  vomers.  The 
two    posterior    parasphenoid    flanges    are 


Fossil  Amiids  •  Boreske        43 


more  splayed  in  A.  jra<^osa  tlian  in  A.  caJva 
or  A.  uintacnsis,  and  overlie  three-fourths 
of  the  basioecipital  length.  As  Estes  ( 1964: 
29)  notes,  there  is  a  relatively  greater  dor- 
soventral  parasphenoid  thiekness  in  A. 
fniiiosa  than  in  A.  calva.  The  parasphenoid 
of  A.  iiiiitoensis  is  proportionately  more 
massive  than  that  of  A.  frai!,osa:  this  mas- 
siveness,  however,  is  probably  a  function 
of  its  greater  size. 

In  A.  fraiiosa,  as  in  A.  uintaensis,  the 
extrascapular  is  tear-drop  shaped,  being 
narrow  at  the  midline  and  expanded  dis- 
tally,  while  in  A.  calva  and  A.  sciitata,  it  is 
more  strap-shaped  and  longer  at  the  mid- 
line. The  proximal  anterior  corner  is 
squared  off,  as  in  A.  scutata  and  A.  calva. 
The  anterior  edge  is  distally  concave  at 
the  pterotic-extrascapular  suture,  and  the 
posterior  edge  is  convex,  particularly 
toward  the  distal  end,  which  is  straight 
rather  than  curved  as  in  the  other  species 
of  Amia.  A.  fragosa  and  A.  uintaensis  lack 
the  posterolateral  projection  displayed  in 
A.  calva. 

The  suprascapular  resembles  that  of  A. 
calva,  except  that  the  distal  edge  is  rela- 
tively straight,  rather  than  incurved.  The 
posterior  border  is  also  straight,  while  in 
A.  calva  there  is  generally  a  slight  concavity 
in  the  middle  of  this  edge;  in  A.  uintaensis 
this  border  is  convex. 

The  pterotic  extends  further  anteriorly 
than  in  A.  calva,  but  not  to  the  extent  that 
it  does  in  A.  uintaensis  or  A.  scutata.  The 
dermosphenotic-pterotic  suture  is  directed 
posterolaterally  in  A.  fragosa  and  antero- 
laterally  in  A.  uintaensis,  A.  scutata,  and 
A.  calva.  As  in  A.  uintaensis  and  A.  scutata, 
this  bone  in  A.  fragosa  is  narrower  an- 
teriorly than  posteriorly,  whereas  in  A. 
calva  the  widths  of  these  ends  are  relatively 
equal. 

The  dermosphenotic  in  A.  fragosa  is  about 
the  same  relative  size  as  in  the  other  species 
of  Amia.  The  anterior  angle  that  forms  the 
posterior  border  of  the  orbit  is  slightly  more 
pronounced  than  in  A.  scutata  and  A.  uin- 
taensis, and  considerably  more  so  than  in 
A.  calva  (Fig.  28). 


The  parietal  in  A.  fragosa  is  character- 
istically square,  whereas  in  A.  calva,  A. 
scutata,  and  A.  uintaensis  it  is  longer  than 
wide.  The  length  of  the  parietal  relative 
to  that  of  the  frontal  is  less  than  in  A.  calva 
and  A.  scutata  and  about  the  same  as  in 
A.  uintaensis.  The  characteristic  deep  ex- 
cavation in  the  frontal  for  the  orbit  is 
displayed  in  all  available  specimens  of 
A.  fragosa.  This  led  Estes  (1964:  36)  to 
postulate  the  presence  of  supraorbital  bones, 
but  the  articulated  specimens  figured  by 
O'Brien  (1969)  show  that  this  was  not  the 
case.  As  Figure  28  shows,  the  ratio  of 
orbital  depth  to  length  is  greater  in  A. 
fragosa  than  in  tlie  other  Amia  species.  As 
noted  in  the  preceding  section  on  the  cranial 
morphometries  of  the  Recent  A.  calva,  it  is 
difficult  to  assign  a  specific  character-state 
of  parietal/frontal  proportions  to  any  of  the 
individual  fossil  Amia  species  because  of 
the  similarity  in  parietal /frontal  propor- 
tions (Table  7).  It  is  apparent,  however, 
that  the  frontals  of  the  earlier  species  A. 
fragosa  and  A.  uintaensis  are  longer  relative 
to  parietal-length  than  in  the  mid-Tertiary 
A.  scutata  or  Recent  A.  calva.  This  feature 
is  useful  in  comparing  A.  fragosa  with  these 
two  species,  but  ineffective  in  distinguishing 
it  from  A.  uintaensis. 

As  Estes  and  Berberian  (1969)  noted, 
the  nasal  displays  a  bifurcation  of  the  an- 
terior border  that  is  lacking  in  A.  calva. 
The  bifurcation  is  also  present  in  A.  uintaen- 
sis, and  the  bone  has  approximately  the 
same  outline  and  size  relative  to  head  size 
as  the  other  forms.  All  available  specimens 
of  A.  fragosa  show  that  the  nasals  lie  much 
closer  to  the  frontals  than  in  A.  calva,  A. 
scutata,  or  A.  uintaensis.  Although  Estes 
( 1964 )  states  that  the  lacrimal  conforms 
closely  with  that  of  A.  calva,  his  restoration 
lacks  the  small  posterior  notch  in  A.  fragosa 
which  accommodates  the  anterior  process 
of  infraorbital  2.  The  lacrimal  in  A.  fragosa 
is  evenly  tapered  at  the  posterior  end,  and 
is  anteroposteriorly  longer  than  in  other 
Amia.  It  is  also  more  dorsoventrally  convex 
than  in  A.  scutata  and  A.  calva. 

As   in   A.   scutata,   infraorbital   4    in   A. 


44         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


fragosa  is  much  more  dorsoventrally  ex- 
panded than  in  A.  calva,  with  the  antero- 
posterior length  extending  almost  to  the 
anterior  edge  of  the  preoperculum.  Infra- 
orbital 4  of  A.  fragosa  and  A.  scutata  is 
more  concave  at  the  dorsal  edge,  and  much 
more  convex  ventrally.  The  pit-line  marks 
extend  further  ventrad  than  is  indicated  in 
the  reconstruction  by  Estes   ( 1964 ) . 

Branchiocranium.  In  A.  fragosa,  the 
supramaxilla  is  relatively  shorter  than  in 
A.  iiintaensis,  A.  scutata,  and  A.  calva,  with 
a  greater  curve  in  the  maxillo-supramaxil- 
lary  suture.  The  dorsoposterior  corner  in 
A.  fragosa  tends  to  be  angular,  as  in  A. 
scutata,  whereas  in  A.  calva  and  A.  uintaen- 
sis  it  is  more  rounded.  The  supramaxilla  is 
deeper  and  more  truncated  at  the  anterior 
end  than  in  other  species  of  Ainia. 

There  is  a  dorsal  shelf  adjacent  to  the 
lingual  border  of  the  alveolar  ridge  which 
widens  the  anterodorsal  surface  of  the  den- 


A.  fragosa  A.  uintaensis      A.  scutata         A.  calva 


A.  fragosa 


A.  uintaensis 


A.   scutata 

Fig.    18.      Comparison      of      mandibles      of 
(transverse    sections    and    ventral    views). 


spp. 


tary  (Fig.  18).  This  shelf  is  lacking  in 
A.  calva  and  A.  uintaensis,  in  which  the 
coronoid  articulation  surface  slopes  directly 
downward  from  the  alveolar  ridge.  This 
region  of  the  lingual  dentary  surface  under- 
lying the  coronoids  extends  more  ventrad 
at  the  symphyseal  edge  than  in  A.  calva, 
and  distinctly  overlaps  the  ventral  part  of 
the  ramus.  There  is  no  such  overlapping  in 
A.  calva;  the  dorsal  and  venti'al  halves  of 
this  region  separate  to  form  Meckel's  groove. 
The  anterodorsal  section  of  the  dentary  in 
A.  uintaensis  overlaps  the  ventral  half,  but 
not  to  the  extent  that  it  does  in  A.  fragosa, 
and  as  the  coronoid  articulation  surface  is 
thicker,  this  thickened  area  of  bone  forms 
the  dorsal  wall  of  Meckel's  groove  as  in 
A.  calva  (Fig.  18).  As  Estes  (1964:  36) 
noted,  the  coronoid  teeth  are  styliform  and 
extend  almost  to  the  ventral  border  of  the 
ramus  at  the  anterior  end;  in  contrast,  the 
coronoid  teeth  of  A.  calva,  A.  scutata,  and 
A.  uintaensis  are  pointed  and  the  coronoids 
do  not  extend  as  far  ventrally  as  in  A. 
fragosa.  The  anterior  half  of  the  dentary 
length  is  more  curved  than  in  A.  calva,  A. 
scutata,  and  A.  uintaensis  (Fig.  18).  This 
is  displayed  in  the  MCZ  9264  specimen 
( Fig.  16 ) ,  in  which  this  curve  approximates 
a  120-degree  angle  at  the  midpoint  of  the 
alveolar  ridge.  The  outline  of  the  dentary 
differs  from  that  of  A.  calva  and  A.  uintaen- 
sis in  that  the  anterior  end  maintains  an 
almost  constant  width  up  to  the  sharp 
curve  at  the  midpoint  of  the  alveolar  ridge, 
at  which  point  it  widens  noticeably.  When 
the  outline  and  curvature  of  the  anterior 
end  of  the  dentary  of  A.  fragosa  are  com- 
pared with  those  of  other  species,  the  result- 
ing difference  appears  to  be  correlated  with 
A.  fragosa's  relatively  smaller  mandible/ 
head  ratio  (Table  7),  smaller  mouth  gape, 
and  its  wider  cranial  roof  (Fig.  15). 

Post-cranial  Skeleton.  On  the  basis  of 
specimens  having  only  the  lateral  surface  of 
the  vertebral  column  exposed,  it  was  con- 
cluded that  centra  of  A.  fragosa  are  indis- 
tinguishable from  those  of  A.  scutata  and 
A.  calva.  Small  disarticulated  vertebrae  are 
also  basically  similar  in  morphology,  there- 


Fossil  Amiids  •  Boreske        45 


fore  it  is  impossible  to  differentiate  A. 
frau^osa,  A.  sciitata,  and  A.  calva.  The  mid- 
trnnk  vertel^rae  of  A.  fra^osa,  A.  sciitata, 
and  A.  calva  differ  from  A.  uintaensis  mid- 
trunk  vertebrae,  which  are  generally  larger, 
and  snbtriangular  rather  than  ovoid.  A. 
fragosa  does,  however,  have  a  vertebral 
column  that  differs  proportionately  and 
meristically  from  that  of  other  species  ( Fig. 
31).  It  has  a  significantly  smaller  total 
number  of  centra  than  the  other  species, 
with  approximately  12  fewer  trunk  verte- 
brae and  8  fewer  monospondylous  caudal 
centra.  It  has  the  same  number  of  diplo- 
spondylous  caudal  centra  as  A.  calva  and 
A.  scutata;  the  number  of  fused  hypurals  is 
also  generally  the  same  (Table  9).  The 
low  number  of  total  vertebrae  in  A.  fragosa 
is  reflected  by  its  shorter,  deeper-bodied 
shape.  The  distance  between  the  anal  fin 
insertion  and  the  end  of  the  vertebral 
column  is  relatively  shorter  than  in  the 
other  species.  The  dorsal  fin  also  terminates 
closer  to  the  caudal  fin  than  in  any  of  the 
other  species  of  Amia  (Plate  1;  Estes  and 
Berberian,  1969:  10).  A.  fragosa  has  fewer 
caudal  lepidotrichia  (19-20)  than  the  other 
species  of  Amia  (23-27).  The  head/ 
standard-length  ratio  of  A.  fragosa  is  greater 
than  that  of  A.  calva,  but  is  not  significantly 
different  from  that  of  A.  uintaensis  or  A. 
scutata  (Table  3).  The  latter  case  is  true 
despite  the  greater  number  of  vertebral 
centra  in  A.  uintaensis  and  A.  scutata;  this 
disparity  may  be  explained  largely  by  the 
fact  that  the  A.  fragosa  skull  itself  is  rela- 
tively shorter  than  that  of  the  other  two 
forms,  particularly  A.  uintaensis,  which  has 
a  greater  head/ standard-length  ratio  than 
A.  fragosa.  Thus  head/ standard-length  does 
not  significantly  reflect  the  length  of  the 
vertebral  column,  but  may  be  used  as  a 
character  with  this  qualification  in  mind. 
The  known  total-length  of  A.  fragosa  falls 
within  the  range  of  A.  calva  and  below  that 
of  A.  uintaensis  (Tables  1-2). 

Discussion 

Marsh  (1871:  105)  described  Amia  new- 
berrianus  and  Amia  depressus  on  the  basis 


of  disarticulated  vertebrae  and  cranial  ele- 
ments from  the  Bridger  Formation  of  Wyo- 
ming. His  main  criteria  for  distinguishing 
these  forms  from  A.  calva  and  from  each 
other  were  that  the  chordal  foramen  of  A. 
newherrianus  was  "considerably  above  the 
center  in  the  dorsal  vertebrae,"  and  that 
A.  depressus  possessed  broader  vertebrae 
than  A.  newherrianus  and  lacked  the  me- 
dian groove  on  the  lower  surface  of  the 
centra.  The  vertebrae  indicated  that  both 
species  were  approximately  the  size  of 
A.  calva.  Osborn  et  al.  (1878:  102)  noted 
that  since  Marsh  gave  no  measurements, 
"the  reference  to  Amia  depressus  cannot  be 
certain."  Marsh  further  noted  that  these 
specimens  belonged  to  the  Yale  College 
Museum,  but  the  specimens  now  seem  to 
have  been  lost.  Marsh  had  apparently  as- 
sumed that  the  characteristics  of  one  verte- 
bra represented  those  of  the  entire  vertebral 
column  and  was  unaware  of  intracolumnar 
variation  in  height/ width  proportions,  aortal 
facet  morphology,  and  position  of  chordal 
foramen  in  the  vertebral  coliunn  of  Amia. 
I  infer  from  Marsh's  report  that  the  type 
specimen  of  A.  depressus  is  probably  a  first 
to  third  trunk  vertebra,  since  the  aortal 
grooves  are  lacking  (Fig.  11)  and  vertebral 
width  exceeds  height  (Fig.  14).  Using  the 
position  of  chordal  foramen  as  a  character 
distinguishing  A.  neivberrianus  is  undiag- 
nostic  since  the  position  of  the  chordal 
foramen  changes  in  relation  to  the  relative 
position  of  the  vertebra  along  the  column 
(Fig.  12).  Therefore,  on  the  basis  of 
Marsh's  undiagnostic  characters  and  the 
similarity  in  size  and  morphology  of  the 
vertebrae  to  those  of  A.  fragosa  and  A. 
calva,  I  consider  both  A.  depressus  and  A. 
newherrianus  as  nomina  duhia. 

Leidy  (1873a:  98)  descrilied  Amia  graci- 
lis from  a  single  trunk  vertebra,  also  from 
the  Bridger  Formation  of  Wyoming.  He 
noted  that  the  centrum  has  two  "oblong 
fossae"  ( aortal  facets )  instead  of  the  charac- 
teristic pair  of  v(>ntral  ridges  found  in  Amia 
calva.  The  size  of  the  centrum  indicated  to 
Leidy  that  A.  gracilis  was  a  smaller  species 
than  A.  calva  (Leidy,  1873b).  The  vertebra 


46         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


(ANSP  5360)  corresponds  approximately 
to  the  twelfth  trunk  vertebra  in  Ainia,  since 
the  aortal  facets  are  oblong  and  unridged 
(Fig.  11).  Although  A.  gracilis  is  small,  it 
falls  well  within  the  size  range  of  A.  fra^osa 
and  A.  calva,  and  is  considered  as  a  nomen 
duhium. 

Estes  and  Berberian  (1969:  10)  suggested 
the  possibility  of  synonymy  of  Paramiatus 
gurleyi  (Plate  1)  and  Atnia  fra<i,osa  with 
the  European  Amia  kehreri  (Plate  2)  on 
the  basis  of  the  close  proximity  of  dorsal 
and  caudal  fins  for  the  former  and  similar- 
ity of  skull  elements  and  teeth  for  the  latter. 
It  was  shown  in  the  previous  section  that 
Paramiatus  <i,urleyi  conforms  not  only  to 
A.  kehreri,  but  also  to  North  American  spec- 
imens of  A.  frafi,osa  on  the  basis  of  body 
morphometries  and  meristics.  Cranial  mor- 
phometries were  also  shown  to  be  similar. 
In  addition,  an  X-ray  (FMNH  X2201)  of 
the  Paramiatus  <^urleyi  skull  reveals  infra- 
orbitals 4  and  5  to  be  longer  than  Romer 
and  Fryxell  (1928)  and  Estes  (1964)  had 
noted.  The  two  infraorbitals  extend  pos- 
teriorly to  the  anterior  edge  of  the  pre- 
operculum  as  they  do  in  A.  frau^osa  (UA 
5398)  from  the  Late  Cretaceous  Edmonton 
Formation  of  Alberta.  The  X-ray  has  also 
revealed  a  displaced  left  vomer  with  26 
styliform  teeth  which  has  been  rotated 
through  the  skull  roof  and  covered  with 
matrix.  All  other  cranial  elements  conform 
morphologically  with  other  Cretaceous  and 
Eocene  specimens  of  A.  frafiosa.  These 
additional  similarities  of  Paramiatus  gurleyi 
and  A.  fragosa  confirm  the  synonymy  of 
these  two  species  which  was  suggested  by 
Estes  and  Berberian  (1969:  10),  and  I 
therefore  include  Paratniatus  gurleyi  in  the 
synonymy  of  Amia  fragosa. 

Comments  on  Related  European  Forms 

Amia  kehreri  was  described  by  Andreae 
(1892,  1895)  from  a  caudal  region,  infra- 
orbital 4,  disarticulated  trunk  vertebrae, 
and  a  left  operculum  from  Middle  Eocene 
(Lutetian)  deposits  at  Messel  bei  Darm- 
stadt    (specimens     at    the     University    of 


Heidelberg,  Andreae  Collection).  On  the 
basis  of  specimens  from  the  same  deposit 
(BMNH  P33480,  Plate  2;  P33488),  it  con- 
forms morphometrically  with  A.  fragosa  in 
head/ standard-length,  pectoral  fin  insertion/ 
standard-length,  mandible/ head-length,  and 
operculum-length/operculum-depth  ( Ta- 
bles 3,  7).  The  distance  between  the  anal 
fin  and  the  end  of  the  vertebral  column 
exceeds  that  of  the  North  American  A. 
fragosa  specimens,  but  is  less  than  in  A. 
scutata  or  A.  calva.  The  parietal/ frontal 
ratio  is  also  greater  than  in  A.  fragosa,  but 
smaller  than  in  A.  scutata  or  A.  calva. 
Meristics  of  vertebral  elements  as  well  as 
the  cranial  characters  discussed  by  Estes 
and  Berberian  ( 1969 )  also  conform  with 
those  of  A.  fragosa.  I  agree  with  Estes  and 
Berberian  ( 1969 :  10 )  that  only  differences 
in  temporal  and  geographical  factors  appear 
to  distinguish  Amia  kehreri  from  A.  fragosa; 
any  osteological  dissimilarities  are  of  minor 
significance. 

The  Middle  Eocene  European  Geiseltal 
deposits  contain  numerous  amiid  fossils; 
according  to  Estes  and  Berberian  ( 1969 ) 
some  showed  resemblances  to  A.  kehreri. 
This  material  is  currently  being  described 
by  Anna  Jerzmanska,  Uniwersytet  Wroclaw- 
ski,  Wroclaw,  Poland. 

Another  related  form  is  Amia  valencien- 
nesi  from  the  Eocene  of  Puy-de-D6me, 
France.  Agassiz  ( 1843 )  described  the  form 
from  one  complete  specimen  and  an  an- 
terior region  of  another  (BMNH  P446, 
27736).  Piton  (1940)  reviewed  these  .speci- 
mens along  with  new  material  collected  at 
the  same  locality.  A.  valenciennesi  also  re- 
sembles A.  kehreri  in  its  vertebral  number 
of  68  centra,  close  approximation  of  dorsal 
and  caudal  fins,  and  an  infraorbital  4  larger 
than  infraorbital  5;  these  similarities  indi- 
cate that  synonymy  with  A.  kehreri  is  in 
order.  The  name  A.  valenciennesi  precedes 
A.  kehreri,  and  thus  has  priority. 

Estes  and  Berberian  (1969:  7)  showed 
that  Amia  russelli  Janot  ( 1966,  1967 )  from 
the  Late  Paleocene  of  France  resembles  A. 
fragosa  in  (!)  square  parietals,  (2)  similar 


Fossil  Amiids  •  Boreske        47 


parietal/frontal  ratio,  (3)  largo  orbital  ex- 
cavation in  frontal,  and  (4)  similar  opc>r- 
cnlnni  height/width  ratio.  Thns  A.  russclli 
conforms  with  many  of  the  most  distinct 
characters  of  A.  kehreri  and  A.  valencien- 
nesi,  and  should  be  considered  a  synonym 
of  the  latter. 

Estes  (1964)  re-evaluated  Dechaseaux's 
( 1937)  redescription  of  the  Early  Oligocene 
Amia  munieri  from  France  and  noted  simi- 
larities with  A.  fraiS,osa  which  included  ( 1 ) 
styliform  vomerine  teeth,  (2)  branchiostegal 
rays  rounded  distally,  ( 3 )  larger  infraorbital 
4  than  infraorbital  5,  and  (4)  similar 
parietal /frontal  proportions.  The  principal 
difference  between  the  forms  is  the  small 
excavation  for  orbits  in  A.  munieri.  Since 
Dechaseaux's  and  Estes'  studies,  the  speci- 
men (MNHN  R4632,  skull  and  associated 
cranial  and  postcranial  elements)  is  being 
further  prepared  to  display  the  cranial  roof 
and  palate  more  extensively.  The  frontal 
lacks  a  prominent  excavation  for  the  orbits 
as  Estes  (1964:  40)  has  noted,  and  in  this 
feature  A.  munieri  resembles  A.  sctitata  and 
A.  calva.  A.  mimieri  is  a  very  important 
form  because  it  represents  the  only  com- 
plete amiid  specimen  known  from  the  Early 
Oligocene,  and,  as  noted,  it  displays  inter- 
mediate morphology  of  the  cranial  features 
among  the  species  of  Amia.  A.  munieri 
occurs  very  late  in  time  in  relation  to  the 
last  known  occurrence  of  A.  jragosa  in 
North  America,  and  because  there  are  no 
complete  specimens  known  from  this  age, 
it  represents  a  stage  of  evolution  among  the 
amiids  that  is  not  found  in  North  America. 

Lehman  ( 1951 )  described  Fseudamia 
lieintzi  (Troms0  Museum  Naturhistorisk 
collections,  Troms0,  Norway)  from  a  fairly 
complete  articulated  sptx-imen  and  two 
skulls  from  probable  Eocene  deposits  in 
Spitzbergen.  He  differentiated  this  form 
from  Amia  on  the  basis  of  ( 1 )  Sinamia-\\ke 
metapterygoid  and  (2)  presence  of  a  con- 
cave notch  on  the  dorsoposterior  border  of 
the  operculum.  Estes  ( 1964 )  noted  that 
Lehman  was  incorrect  in  his  interpretation 
of  tlie  nature  of  the  metapterygoid  and 
operculum,    and    therefore    suggested   that 


Fseudamia  might  be  placed  in  the  genus 
Amia.  From  the  examination  of  Lehman's 
plates,  it  appears  that  this  form  resembles 
A.  fra^osa  in  its  deep-bodied  shape  and 
low  parietal /frontal  ratio  (approximately 
0.410),  and  that  it  may  be  synonymous  with 
A.  valenciennesi  and  A.  kehreri.  I'urther 
preparation  would  possibly  be  helpful  in 
uncovering  palatal  teeth,  whose  moiphology 
would  aid  in  a  more  definitive  description. 
Although  the  exact  age  of  the  Eocene 
deposit  in  which  the  specimen  occurred 
is  uncertain,  this  Spitzbergcni  locality,  if 
Early  Eocene,  lies  on  the  possible  migration 
route  of  amiids  (and  other  vertebrates) 
between  North  America  and  Europe. 

Amia  uintaensis  (Leidy,  1873) 

Protamia  tiintacnsis  Leidy,  1873a:   98. 
Protamia  media  Leidy,  1873a:   98. 
Pappichthy.s  plicatus  Cope,  1873:  635. 
Pappichthtjs  sclerops  Cope,  1873:  635. 
Pappichthy.s  laevis  Cope,  1873:  636. 
Pappichthijs  symphysis  Cope,  1873:  636. 
Pappichthys  corsonii  Cope,  1873:   636. 
Pappichthys  meditis  Cope,  1884:  pi.  4. 
Amia  ivhiteavcsiana  Cope,  1891:  2. 
Amia  macrospondyla  Cope,  1891:  2. 

Holotype.  ANSP  5558,  anterior  tiimk 
vertebra. 

Paratypes.  ANSP  8044,  first  anterior 
trunk  vertebra;  ANSP  3151,  three  posterior 
trimk  vertebrae;  ANSP  5622,  basioccipital. 

Type  locality  and  horizon.  Henrv's  Fork. 
North  half  of  section  5,  T  12  N,  R  111  W, 
Sweetwater  County,  Wyoming;  Bridger 
Formation. 

Age  rouge.  Torrejonian  (Middle  Paleo- 
cene)  to  Chadronian  (Early  Oligocene). 

Hypodi^m.  Paleocene.  Fort  Union  For- 
mation, Wyoming  and  Montana:  PU  17117, 
maxillary;  PU  17068,  vertebrae  and  denta- 
ries;  PU  162.36,  disarticulated  skull  and 
trunk  vertebrae;  CM  25364,  dentary;  PU 
17064,  trunk  vertebrae.  Tongue  River  For- 
mation, Montana:  PU  20578,  basioccipital 
and  vertebrae.  Paskapoo  Formation,  Al- 
berta: ROM  4653,  vertebrae. 

Eocene.  Will  wood  Formation,  Wyo- 
ming: PU  21173,  basioccipitals;  PU  17227, 
basioccipital  and  trunk  vertebrae;  PU  17649, 


1 


48         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


portion  of  cranium;  PU  18760,  skull  frag- 
ments, dentary,  and  vertebrae.  Wasatch 
Formation,  Wyoming:  AMNH  4635,  dentaiy 
and  maxilla.  Golden  Valley  Formation, 
North  Dakota:  PU  18568,  basioccipital. 
Green  River  Formation,  Wyoming:  USNM 
18147,  skull  fragments  and  vertebrae; 
AMNH  785,  complete  caudal  region;  PU 
13865,  nearly  complete  specimen;  MCZ 
12916,  disarticulated  skull  and  associated 
vertebrae.  Wind  River  Formation,  Wyo- 
ming: AMNH  2437,  dentary  and  skull  frag- 
ments. Bridger  Formation,  Wyoming:  CM 
25362,  portion  of  cranium  and  vertebral 
column;  AMNH  4631,  portion  of  cranium 
with  dentaries,  gular,  and  basioccipital; 
USNM  170976,  maxilla;  YPM  6238-6240, 
6242,  6244,  6250-6253,  6257-6258,  vertebrae 
and  basioccipitals;  USNM  170973,  5450, 
3962,  3963,  3966,  PU  20523,  10101,  ANSP 
2337-2339,  vertebrae;  USNM  2181,  ANSP 
5632,  trunk  vertebrae;  USNM  3959,  trunk 
and  caudal  vertebrae;  ANSP  5580,  mid- 
trunk  vertebra;  AMNH  2539,  anterior  por- 
tion of  a  left  dentary,  two  premaxillae,  right 
quadrate,  left  epihyal,  anterior  portion  of  an 
ectopterygoid,  three  trunk  vertebrae,  and 
numerous  fragments  of  angular;  USNM 
3965,  left  dentary;  USNM  3968,  anterior 
dentary  fragment;  AMNH  2570,  pre-maxil- 
lary  fragment,  fragments  of  angular,  left 
quadrate  fragment,  trunk  vertebra  frag- 
ment, and  a  caudal  vertebra;  USNM  3960, 
PU  10099,  10110,  vertebrae  and  a  ural  cen- 
trum; USNM  5476,  basioccipital;  USNM 
3961,  left  dentary  fragment.  Washakie  For- 
mation, Wyoming:  FMNH  27465,  4509,  ver- 
tebrae. Uinta  Formation,  Utah:  CM  2382, 
maxillary  fragment. 

Oligocene.  Cypress  Hills  Formation, 
Saskatchewan:  NMC  6197,  trunk  vertebra; 
NMC  6198,  caudal  vertebra. 

Known  distribution.  Montana,  Wyo- 
ming, Utah,  North  Dakota,  Alberta,  and 
Saskatchewan. 

Revised  diagnosis.  Vertebral  column 
with  approximately  20  more  vertebral  seg- 
ments in  total  number  (85)  than  A.  fragosa, 
and  five  fewer  trunk  centra  (31)  and  five 
more     diplospondylous     caudal     vertebrae 


(21)  than  in  the  other  long-bodied  forms, 
A.  scutata  and  A.  calvo.  Mid-trunk  verte- 
brae subtriangular  rather  than  ovoid.  Pa- 
latal teeth  sharp,  greatly  curved  inwardly. 
Between  40-45  vomerine  teeth  as  compared 
with  15-17  in  A.  fragosa,  A.  scutata,  and 
A.  calva.  Hyomandibular  more  deeply 
notched  between  opercular  process  and 
extensor  (dorsal)  surface  than  in  other 
species;  opercular  process  relatively  larger. 
Angle  between  alveolar  ridge  and  exterior 
surface  of  the  dentary  forms  a  more  acute 
angle  than  in  the  other  species.  Mandibular 
ramus  less  curved  than  in  other  species,  so 
that  angle  between  symphyseal  ends  of 
dentaries  is  relatively  narrow.  Greater 
mandible/head  ratio  (0.693)  and  head/ 
standard-length  ratio  (0.322)  than  any  of 
the  other  forms:  A.  uintaensis  has  a  head 
relatively  longer  and  a  mouth  gape  rela- 
tively wider  than  do  other  species.  Most 
specimens  are  significantly  larger  than  the 
other  species,  with  a  relatively  greater  de- 
gree of  ossification  of  all  bones.  Greatest 
known  standard-length  800  mm. 

Introduction 

Leidy  (1873a)  reported  numerous  dis- 
articulated vertebrae  of  a  fossil  fish  related 
to  Amia  from  the  Bridger  Formation  of 
Wyoming.  He  distinguished  a  new  genus 
Protamia  from  Amia  by  its  "two  oval  fossae" 
( aortal  facets )  on  the  ventral  surface  of  the 
centrum,  and  by  large  vertebrae  character- 
istically with  a  much  greater  width  to 
height  proportion.  Hijpamia,  another  new 
genus  from  the  same  locality  which  Leidy 
also  related  to  Amia,  was  characterized  by 
also  being  larger  than  A.  calva,  and  by 
vertebrae  whose  sides  converged  into  a 
"medium  prominence  excavated  into  a  pair 
of  oval  fossae"  deeper  than  those  of  Pro- 
tamia. Later  ( 1873b ) ,  Leidy  published  a 
more  complete  and  illustrated  account  of 
the  various  species  of  the  new  genera 
Protamia  and  Hypamia.  In  the  same  year 
Cope  ( 1873 )  described  a  new  amiid  genus, 
also  from  the  Bridger  Formation,  which  he 
named  Pappichthys.  He  distinguished  this 
new  genus  from  Amia  by  the  "presence  of 


Fossil  Amiids  •  Boreske 


49 


only  one  series  of  teeth,  instead  of  several, 
on  the  bones  about  the  mouth."  Osborn 
et  al.  ( 1878 )  reported  other  finds  of  Pci})- 
picJitJiys  from  the  Bridger  Formation  which 
seemed  to  fit  Cope's  description.  Cope 
(1884)  further  discussed  his  new  genus, 
and  rejected  Leidy's  prior  nomenclatiue 
and  description. 

New  ton  ( 1899 )  discussed  this  nomencla- 
tural  controversy  and  asserted  the  validity 
of  Leidy's  genius  Protamia,  since  Cope's 
later  diagnosis  \\'as  no  more  effective  in 
characterizing  the  new  genus  than  Leidy's 
prior  one.  Newton  bc^lieved  that  Cope's 
description  of  PappicJitlitjs  as  having  only 
a  single  row  of  marginal  teeth  was  taxo- 
nomically  undiagnostic,  since  this  condition 
would  also  include  A.  calva.  Romer  and 
Fryxell  ( 1928 )  accepted  Leidy's  earlier 
description  and  genus  as  diagnostic,  and 
referred  PappiclitJujs  to  Protamia.  They 
also  mentioned  Hypamia  but  found  little 
to  distinguish  it  from  Amia. 

Hussakof  (1932)  continued  to  use  Cope's 
name,  however,  and  reported  large  speci- 
mens of  Pappichthys  from  the  Eocene  of 
Mongolia.  He  also  noted  Cope's  error  in 
diagnosing  the  tooth  characteristics  of  the 
genus,  since  Pappichthys  {Protamia)  has 
several  rows  of  small  teeth  on  the  "splenial 
bone."  In  comparison  with  Amia  he  noted 
"points  of  difference  in  nearly  every  bone 
available  for  comparison,"  and  concluded 
that  Pappichthys  was  a  valid  genus,  "not 
merely  a  group  of  large-sized  extinct  species 
of  Amiatus." 

Estes  (1964),  like  Romer  and  Fryxell 
(1928),  referred  Pappichthys  to  Protamia, 
and  reported  several  vertebrae  and  a  maxil- 
lary fragment  from  the  Cretaceous  Lance 
Formation  of  Wyoming.  He  inteipreted  the 
increase  in  breadth  over  thickness  of  the 
vertebrae  as  a  po.ssible  "function  of  in- 
creased size,"  a  condition  that  would  also 
allow  for  tlie  comparatively  more  massive 
nature  of  the  maxillary  fragment.  He  also 
considered  the  retention  of  this  genus  as 
arbitrary  until  enough  materials  were  avail- 
able. Janot  (1967)  did  not  consider  this 
single   distinguishing   characteristic   of   the 


vertebrae  as  sufficient  foundation  for  the 
erecti(m  of  a  new  genus,  and  therefore  sug- 
gested relerring  Protamia  to  Amia.  Estes 
et  al.  (1969)  concurred  with  Janot  in 
synonymizing  Protamia  with  Amia.  The 
present  study  confirms  tliis  synonymy; 
Leidy's  species  (1873a)  has  priority  and 
the  valid  name  of  this  fish  is  thus  the  oldest 
specific  name,  Amia  uintacnsis. 

Revision  of  all  forms  referred  now  or  in 
the  past  to  Protamia  is  much  needed,  for 
these  large  amiids  were  diagnosed  on  char- 
acters of  isohited  vertebrae  and  skull  frag- 
ments. This  study  gives  more  useful 
diagnostic  characters  that  provide  a  basis 
on  which  the  taxonomy  of  this  group  can  be 
established. 

Fossil  Record 

The  major  deposits  carrying  remains  of 
Amia  uintacnsis  (Table  18)  range  in  age 
from  Middle  Paleocene  to  Early  Oligocene. 
Middle  Paleocene  specimens  occur  in  the 
Fort  Union,  Tongue  River,  and  Paskapoo 
formations  and  consist  mostly  of  isolated 
and  broken  centra,  and  dentary  and  maxil- 
lary fragments.  A  nearly  complete  skull 
(PU  162.36)  with  associated  trunk  and 
caudal  centra  from  the  Bear  Creek  local 
fauna  of  Montana  (Fort  Union  Formation) 
is  the  only  articulated  specimen  from  the 
Late  Paleocene.  The  Eocene  material  in- 
cludes one  complete  articulated  specimen 
(PU  13865),  one  complete  caudal  region 
(AMNH  785),  and  a  disarticulated  skull 
(MCZ  12916)  from  the  Creen  River  Forma- 
tion. PU  13865  (Plate  3)  has  the  axial 
skeleton  intact  in  matrix,  with  a  dislocated 
fifth  centrum  that  is  the  only  one  available 
for  three-dimensional  measurements.  This 
is  also  the  only  specimen  in  which  a  com- 
plete vertebral  count  can  be  taken.  AMNH 
785  provides  excellent  meristic  information 
for  the  caudal  region  (Fig.  8C).  CM  25362, 
from  the  Bridger  Formation,  consists  of  a 
left  palatal  and  opercular  series  and  an 
almost  complete,  disarticulated  vertebral 
column  that  permitted  the  taking  of  a  series 
of  centrum  measurements.  Other  skull  frag- 
ments and  vertebrae  occur  in  many  deposits 


50         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


throughout  the  Eocene  (Table  18).  The 
latest  occurrence  of  A.  uintaensis  is  repre- 
sented by  two  isolated  centra  from  the 
Cypress  Hills  Formation  (Oligocene,  Chad- 
ronian ) . 

Description 

Neurocranhan.  Posterior  to  the  spinal 
arterial  foramina  the  basioccipital  includes 
two  fused  vertebrae.  As  the  basioccipitals 
display  great  variation  in  the  morphology  of 
the  articular  surface,  it  is  difficult  to  char- 
acterize this  form  on  the  basis  of  this 
feature.  However,  the  articular  surface  is 
generally  kidney-shaped,  with  dorsal  in- 
dentations bet\veen  the  neural  facets,  and 
ventrally  there  is  an  indentation  distal  to 
the  aortal  facets.  In  A.  froii^osa  and  A. 
calva  the  basioccipital  has  ovoid  articular 
surfaces  with  no  dorsal  indentations  be- 
tween the  neural  facets  (Estes,  1964:  29, 
fig.  15).  In  lateral  view  the  distal  articular 
surface  of  the  A.  uintaensis  basioccipital  is 
not  perpendicular  to  the  parasphenoid 
flanges;  the  dorsal  half  of  this  surface  is 
more  anteriorly  directed  than  the  ventral 
half. 

The  parasphenoid  is  longer  relative  to 
its  width  than  it  is  in  either  A.  calva  or 
A.  fragosa,  primarily  in  the  region  anterior 
to  the  ascending  processes.  At  the  point 
nearest  the  ascending  processes,  it  lacks 
the  pronounced  convexity  and  die  accom- 
panying anterior  lateral  notches  found  in 
A.  calva  and  A.  fragosa.  The  ascending 
processes  are  slightly  less  anteriorly  oriented 
in  ventral  view  than  in  A.  calva,  but  more 
so  than  in  A.  fragosa  ( Fig.  17 ) .  The  region 
posterior  to  the  ascending  processes  is  rela- 
tively shorter  than  in  A.  fragosa  or  A.  calva; 
it  is  also  more  massive  and  more  ventrally 
convex  than  in  the  other  two  forms.  The 
posterior  parasphenoid  flanges  resemble 
tliose  of  A.  calva  more  tlian  A.  fragosa  in 
outline  as  well  as  juxtaposition;  those  of 
A.  fragosa  are  more  laterally  splayed  than 
in  A.  uintaensis  or  A.  calva.  The  tooth-bear- 
ing surface  differs  considerably  from  that  of 
A.  fragosa  and  somewhat  from  A.  calva  in 
outline   and   extent.    As   in   A.    calva,   this 


surface  extends  anteriorly  to  the  vomers, 
but  its  width  is  much  greater  and  more 
constant  than  in  A.  calva,  which  is  narrowly 
tapered  anteriorly.  Posteriorly,  this  surface 
extends  further  than  in  A.  calva,  but  not  as 
far  as  in  A.  fragosa.  Approximately  two- 
thirds  of  the  tooth-bearing  surface  lies 
anterior  to  the  ascending  processes,  while  in 
A.  calva  this  area  is  anteroposteriorly  cen- 
tered, and  in  A.  fragosa  it  is  nearly  all 
posterior.  The  tooth-bearing  surface  covers 
a  greater  portion  of  the  ventral  surface  of 
the  parasphenoid  than  in  A.  fragosa  or  A. 
calva;  its  basic  outline  is  diamond-shaped, 
with  the  anterior  apex  widened  and  ex- 
tended to  the  vomers,  while  that  of  A. 
fragosa  is  subrectangular  and  that  of  A. 
calva  is  tear-drop  shaped  with  the  apex 
sharply  protracted  anteriorly. 

In  A.  uintaensis  the  distal  edge  of  the 
suprascapular  is  convex  as  in  A.  calva,  while 
in  A.  fragosa  this  edge  is  almost  a  straight 
line.  The  posterior  border  is  more  rounded 
distally  than  in  A.  calva  and  is  convex  rather 
than  concave. 

In  having  the  extrascapular  rounded  at 
the  distal  border,  A.  uintaensis  is  the  same 
as  A.  calva  and  A.  scutata,  but  differs  from 
both  of  them  in  that  th(^  posterior  border 
is  not  concave,  and  from  A.  calva  alone  in 
lacking  the  distal  posterior  process.  The 
anterior  border  is  relatively  straight,  unlike 
the  condition  in  A.  calva  and  A.  scutata,  in 
which  the  lateral  distal  ends  of  the  anterior 
borders  are  directly  posteriad.  As  in  A. 
fragosa  the  midline  is  shorter  than  in  A. 
scutata  and  A.  calva. 

As  in  A.  scutata  and  in  A.  fragosa  the 
pterotic  is  narrower  at  the  anterior  than 
posterior  border,  while  in  A.  calva  and,  to 
an  extent,  in  A.  scutata  the  ends  are  sub- 
equal.  As  in  A.  fragosa  they  extend  farther 
anteriorly  and  adjoin  the  frontal s  postero- 
laterally.  The  dermosphenotic-pterotic  su- 
ture is  anterolaterally  directed,  as  in  A. 
scutata,  but  not  as  pronoimced  as  in  A. 
calva.  The  anterolateral  edge  of  the  pterotic 
is  indented  and  forms,  witli  the  dermo- 
sphcnotic,  an  additional  concavity  in  the 
outline  of  the  cranial  roof.   Aside  from  this 


Fossil  Amiids  •  Boreske        51 


anterior  indentation,  the  lateral  borders  are 
relatively  straight,  as  eompared  with  the 
smoothly  coneave  exterior  sides  of  the 
pteroties  in  A.  scututa,  A.  calva,  and  A. 
fruf!,osa.  The  posterior  border  forms  a 
smooth  line,  as  in  A.  fragosa,  and  laeks  the 
small  lappet  that  A.  scutata  and  A.  calva 
display. 

The  dermosphenotie  is  similar  to  that  ot 
A.  calva  in  relative  size  and  outline,  al- 
though it  does  not  jut  as  deeply  into  the 
frontals.  Its  anterior  border  is  rounded,  as 
in  A.  calva,  rather  than  sharply  angular,  as 
in  A.  fra^osa.  The  posterior  half  of  the 
outer  lateral  border  is  indented  to  form  a 
coneavity  with  the  anterior  tip  of  the 
pteroties.  The  parietal  in  A.  uintaensis  is 
elongated  anteriorly,  as  in  A.  calva  and  A. 
scutata,  while  that  of  A.  jra^osa  is  relatively 
square.  The  orbital  excavation  in  the  lateral 
sides  of  the  frontal  is  shallow  as  in  A.  calva 
and  A.  scutata,  while  that  of  A.  fragosa  is 
characteristically  deep  (Fig.  28).  The  sen- 
sory canal  cannot  be  determined.  The 
frontals  are  more  elongated  relative  to 
parietal  length  tlian  in  A.  calva  and  A. 
scutata;  the  parietal /frontal  ratio  is  only 
slightly  smaller  than  that  of  A.  fragosa 
(Table  7).  The  distal  lateral  border  tapers 
anteromedially,  and  the  anterior  ends  are 
relatively  pointed  anteriorly,  forming  a  deep 
notch  on  the  midline  suture. 

There  is  a  slight  bifurcation  of  the 
anterior  border  of  the  nasal  as  in  A.  fragosa. 
The  nasal  bones  are  relatively  narrower 
than  in  A.  fragosa  or  A.  calva,  but  are 
otherwise  similar  in  shape  and  relative  size. 
They  are  fairly  well  separated  from  the 
frontals,  as  in  A.  calva  and  A.  scutata,  rather 
than  abutting  them  as  in  A.  fragosa. 

The  lacrimal  in  A.  uintaensis  resembles 
that  of  A.  fragosa  in  general  morphology, 
although  it  lacks  the  posterior  notch  for  the 
anterior  end  of  infraorbital  2  which  is 
present  in  the  other  species  of  Amia.  The 
lacrimal,  like  that  in  A.  fragosa,  is  relatively 
longer  and  more  tapered  posteriorly  than  in 
A.  scutata  and  A.  calva.  It  is  more  dorsally 
convex  than  in  the  other  forms,  but  only 
slightly  more  so  than  in  A.  fragosa. 


The  infraorbital  5  in  A.  uintai'nsis  is 
similar  to  that  in  A.  fragosa  and  A.  scutata, 
being  less  robust  posteriorly  than  in  A. 
calva.  As  in  the  other  forms,  it  is  narrower 
anteriorly  than  posteriorly.  The  ventral 
border  is  relatively  straight,  while  that  of 
the  other  forms  is  posteriorly  convex.  Infra- 
orbitals 2,  3,  and  4  have  not  been  identified. 

The  vomerine  tooth  patch  in  A.  uintaen- 
sis, as  in  A.  fragosa,  extends  inore  posteri- 
orly than  in  A.  calva  (Fig.  19).  The 
\^omerine  teeth  are  sharp  and  greatly  curved 
posteriorly;  they  exceed  those  of  A.  fragosa 
and  A.  calva  in  number,  each  vomer  bear- 
ing between  40-50  teeth,  as  compared  to 
half  that  number  in  A.  fragosa  and  A.  calva. 
The  rostral  and  antorbital  are  identical  to 
that  of  the  other  species. 

Brancliiocranium.  The  suture  between 
the  anterior  and  posterior  dermopalatine 
cannot  be  discerned.  In  A.  uintaensis  the 
dermopalatine  has  about  twice  the  number 
of  teeth  as  in  A.  calva,  and  the  tooth  patch 
extends  more  distad.  The  teeth  are  sharply 
pointed,  as  are  the  vomerine  teeth. 

The  hyomandibular  is  more  deeply  ex- 
cavated between  the  opercular  process  and 
the  extensor  ( dorsal )  surface,  and  the  oper- 
cular process  is  more  massive  and  extends 
further  ventrad,  forming  a  larger  articula- 
tion surface,  as  compared  with  the  other 
species  of  Amia.  The  articular  surface  of 
the  quadrate  is  more  robust  than  in  other 
species  of  Amia  and  displays  three  cristae 
ventrally  rather  tlian  dorsally  as  in  A.  calva 
and  A.  rolmsta  (Janot,  1967:  144).  The 
ceratohyal  resembles  that  of  A.  calva  and 
A.  fragosa  with  the  exception  of  its  being 
thicker  at  the  neck  of  the  proximal  end. 
The  metapterygoid  in  A.  uintaensis  con- 
forms very  closely  to  that  of  A.  calva  in 
outline  and  in  the  position  of  the  anterior 


Fig.    19.      Comparison  of  vomers  of  A,  Amia  calva;  B,  A. 
uintaensis;  and  C,  A.  fragosa. 


52        Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  1 


basal  process  and  the  posterolateral  otic 
process. 

The  maxilla  in  A.  uintaensis  is  more 
robust  and  relatively  longer,  and  its  pos- 
terior border  is  dorsoventrally  wider  than 
in  the  other  forms,  particularly  A.  fragosa. 
As  in  A.  calva  the  small  supramaxillary 
notch  occurs  more  anteriorly  than  in  A. 
fragoso.  The  dorsoposterior  border  is 
rounded,  as  in  A.  calva  and  A.  scutata, 
rather  than  sharply  angular,  as  in  A.  fragosa. 
Anteriorly  the  maxilla  is  deeper  and  more 
thickly  ossified  than  in  the  other  forms, 
but  this  may  be  a  function  of  greater  size. 
The  supramaxilla  resembles  that  of  A.  calva 
in  general  morphology,  being  elongated  and 
narrowly  tapered  anteriorly,  with  a  smooth- 
ly rounded  posterior  end  conforming  to  the 
curve  of  the  maxilla.  The  maxillo-supra- 
maxillary  suture  is  straight  as  in  A.  calva. 
The  premaxilla  is  identical  to  that  of  the 
other  species. 

The  dentary  of  A.  uintaensis  is  similar  to 
that  of  A.  calva  and  A.  scutata  in  lacking 
the  dorsal  shelf  of  the  anterior  lingual  bor- 
der of  the  alveolar  ridge  which  occurs  in 
A.  fragosa.  The  coronoids  articulate  more 
or  less  vertically  on  the  alveolar  ridge,  as  in 
A.  scutata  and  A.  calva.  The  anterodorsal 
region  of  the  dentary  slightly  overlaps  the 
ventral  half,  but  not  to  the  extent  that  it 
does  in  A.  fragosa;  A.  uintaensis  seems  to 
be  intermediate  between  A.  fragosa  and  A. 
calva  in  this  feature,  the  latter  having  no 
such  ventral  overlapping  at  the  symphyseal 
edge.  The  coronoid  articulation  surface  of 
the  A.  uintaensis  dentary  is  thicker  than  in 
A.  fragosa  and  A.  calva,  but  only  slightly 
more  so  than  in  A.  scutata.  At  the  termina- 
tion of  this  surface,  this  thickened  area  of 
bone  forms  the  dorsal  wall  of  the  Meckelian 
groove,  as  in  A.  calva.  The  ventral  wall  of 
this  groove  is  less  well  defined  than  in  A. 
calva,  witli  A.  scutata  being  intermediate. 
The  anterior  half  of  the  dentary  length  in 
A.  uintaensis  is  evenly  tapered  to  the 
symphyseal  edge;  it  is  elongated  and  lacks 
the  sharp  curve  present  in  A.  fragosa  at  the 
midpoint  of  the  alveolar  ridge  (Fig.  18). 
There  is  only  a  trace  of  such  a  curve  in  the 


dentaries  of  A.  calva  and  A.  scutata  which 
are  also  more  elongated  and  evenly  tapered 
than  in  A.  fragosa,  although  not  to  the  ex- 
tent that  they  are  in  A.  uintaensis.  Anteri- 
orly, the  bone  is  also  relatively  thicker  than 
in  A.  fragosa  and  A.  calva;  A.  scutata  also 
displays  this  greater  ossification  at  the 
anterior  end  of  the  dentary.  Posteriorly, 
the  dentary  is  very  similar  to  that  of  A. 
calva.  The  coronoid  teeth  are  sharp  and 
conelike,  extending  to  the  midpoint  of  the 
lingual  surface,  as  in  A.  calva.  As  Janot 
(1967)  shows  for  A.  robusta,  the  alveolar 
ridge  is  more  horizontal  in  A.  uintaensis  and 
forms  a  more  acute  angle  with  the  exterior 
surface  of  the  dentary  than  it  does  in  A. 
fragosa  or  A.  calva;  A.  scutata  is  interme- 
diate between  A.  uintaensis  and  A.  calva  in 
this  feature  (Fig.  18).  In  A.  uintaensis  the 
first  coronoid  (symphyseal)  overlies  only 
the  dorsal  half  of  the  anterior  articular  sur- 
face of  the  dentary,  as  in  A.  calva  and  A. 
scutata.  The  teeth  are  more  sharply  pointed 
than  in  any  of  the  other  forms  (Fig.  18). 
The  second  coronoid  is  fragmentary,  but 
appears  to  resemble  that  of  A.  calva  with 
the  exception  of  its  having  more  sharply 
pointed  teeth.  The  prearticular  specimens 
available  are  fragmentary,  but  the  lingual 
surface  possesses  blunt-conical  teeth  similar 
to  those  in  A.  calva  and  A.  fragosa.  Bor- 
sally,  however,  these  teeth  are  as  sharply 
pointed  as  the  coronoid  teeth.  The  angular 
is  slightly  longer  and  higher  than  that  of 
A.  calva.  The  posterior  border  is  more  ver- 
tical, with  the  articular  notch  less  pro- 
nounced. It  is  more  heavily  ossified  than 
in  A.  calva,  but  this  may  be  a  function  of 
size.  The  surangular  in  A.  uintaensis  is 
basically  similar  to  that  of  A.  calva,  al- 
though it  is  situated  more  dorsally  and  is 
more  rounded  at  the  dorsal  edge. 

The  gular  is  longer  than  that  of  A.  calva 
and  A.  fragosa  (Fig.  20).  It  is  also  slightly 
narrower  at  the  posterior  end  than  the 
anterior  end,  while  the  reverse  is  generally 
true  in  A.  calva.  Otherwise,  the  gular 
strongly  resembles  that  of  A.  calva.  Despite 
a  few  minor  dissimilarities,  the  preopercu- 
lum  resembles  that  of  A.  calva.   There  is  a 


Fossil  Amiids  •  Boreske        53 


Fig.   20.      Comparison    of    gulars    of    A,    Amia    calva;    B, 
A.    uintaensis;    and    C,    A.    fragosa. 


slightly  more  pronounced  concavity  in  the 
ventroposterior  border  than  is  exhibited  in 
A.  calva;  this  concavity  is  altogether  lacking 
in  A.  fragosa.  The  line  of  curvature  is  about 
the  same  as  in  A.  calva;  in  A.  fragosa  the 
preoperculum  is  more  deeply  curved.  The 
dorsal  half  is  not  quite  as  wide  as  the 
ventral  half,  while  in  A.  calva  both  ends  are 
fairly  equal.  In  A.  fragosa,  however,  the 
dorsal  half  is  much  narrower  and  more 
tapered  than  the  ventral  half,  which  is  rela- 
tively wider  and  bulbous.  The  operculum 
in  A.  uintaensis  is  similar  to  that  of  A. 
calva  and  A.  scutata  in  operculum-depth/ 
operculum-length  (Table  7).  The  suboper- 
culum  conforms  in  general  morphology  with 
that  of  A.  calva,  although  it  is  slightly  more 
robust,  particularly  in  the  posterior  region. 
The  corners  tend  to  be  angular,  as  in  A. 
scutata  and  A.  calva,  rather  than  rounded, 
as  in  A.  fragosa.  The  interoperculum  is 
similar  to  that  of  A.  calva,  although  more 
robust.  The  anterodorsal  border  is  more 
convex  than  in  A.  calva,  and  is  more  deeply 
impressed  into  the  preoperculum.  The 
anteroventral  border  is  narrowly  tapered  as 
in  A.  calva,  rather  than  smoothly  rounded  as 
in  A.  fragosa.  The  first  branchiostegal  ray 
conforms  to  that  of  the  other  species.  Al- 
though the  lack  of  articulated  material 
makes  any  count  of  the  rays  difficult,  in 
MCZ  12916  there  are  12  disarticulated 
branchiostegal  rays  on  the  right  side  of  the 
cranial  roof.  As  in  A.  fragosa  the  distal  ends 
of  the  rays  are  consistently  rounded,  rather 
than  squared  as  in  A.  calva. 

Post-cranial  skeleton.  The  supracleithrum 


in  A.  uintaensis  resembles  that  of  A.  calva 
and  A.  fragosa,  excepting  the  dorsal  articu- 
lation surface,  which  is  rectilinear  rather 
than  pointed  as  in  A.  calva.  The  distal 
lateral  border  in  the  Paleocene  specimens 
lacks  the  notch  that  occurs  in  A.  calva,  but 
this  notch  is  present  in  the  Eocene  speci- 
mens. The  metacleithrum  in  A.  uintaensis 
is  more  elongated  than  in  A.  calva  and  A. 
fragosa.  The  dorsal  end  is  narrower  than 
in  A.  calva,  and  the  ventral  end  is  sciuared 
off.  The  cleithrum  in  A.  uintaensis  is  largely 
similar  to  that  of  the  other  Ainia  species, 
but  is  more  massive  at  the  proximal  end 
than  in  A.  calva,  and  the  dermal  sculpture 
covers  a  greater  area  than  in  A.  calva,  ex- 
tending to  the  distal  border  as  in  A.  fragosa 
and  A.  scutata  (Fig.  21).  The  mid-distal 
border  is  smoothly  convex  and  lacks  the 
notch  ventral  to  the  metacleithrum  which 
is  present  in  A.  calva. 

The  preceding  study  of  the  vertebral 
skeleton  of  A.  calva  revealed  changes  in 
height/ width  proportions,  position  of 
chordal  foramen,  configuration  of  neural 
and  aortal  facets,  and  in  the  basapophyscal 
angles  and  length  of  basapophyses  which 
may  be  used  here  to  discern  similar  trends 
in  A.  uintaensis  centra,  for  the  fossil  verte- 
brae display  the  same  features  characteris- 
tic of  the  Recent  species  even  in  disarticu- 
lated state. 

CM  25362  from  the  Bridger  Formation  is 
the  only  specimen  that  has  a  relatively 
complete,  disarticulated,  undistorted  verte- 


B 


Fig.   21.      Comparison    of    cleithra    of    A,    Amia    calva;    B, 
A.   scufata;   C,   A.    uintaensis;   and    D,   A.    fragosa. 


54         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


bral  column;  as  the  centra  are  separable  this 
specimen  is  useful  in  comparisons  with  iso- 
lated vertebrae.  There  are  59  centra  pres- 
ent: 25  trunk  centra  and  34  caudal  centra, 
including  two  fused  urals.  Many  of  the 
preserved  caudal  centra  are  only  fragments. 
Since  the  articulated  specimen  (PU  13865) 
has  85  vertebrae  (see  Table  9  for  regional 
numbers)  it  may  be  assumed  that  about  25 
vertebrae  are  missing  from  CM  25362. 
When  comparing  vertebrae  from  different 
regions  of  the  column  in  the  two  specimens, 
it  appears  that  CM  25362  lacks  approxi- 
mately six  trunk  and  approximately  twenty 
caudal  centra.  The  first  anterior  trunk  cen- 
trum present  in  the  CM  25362  series  pos- 
sesses aortal  facet  configurations  similar  to 
those  of  the  seventh  vertebra  of  the  articu- 
lated specimen  ( PU  13865 ) .  An  articulated 
but  separable  series  of  six  uncrushed  an- 
terior trunk  vertebrae  (PU  10101),  also 
from  the  Bridger  Formation,  aids  in  the 
reconstruction  of  the  anterior  region  of  the 


A.  uintaensis  vertebral  column  (Figs.  22- 
25).  The  basapophyseal  angles  of  these  six 
PU  10101  vertebrae  do  not  vary  from  180 
degrees.  The  first  six  anterior  trunk  verte- 
brae from  a  partly  disarticulated  vertebral 
column  from  the  Paleocene  specimen  (PU 
16236)  resemble  the  six  PU  10101  centra  in 
length  and  shape  of  aortal  facets,  even 
though  PU  16236  is  a  smaller  individual. 
The  height/ width  ratio  of  these  latter  centra 
is  difficult  to  determine,  however,  since  the 
specimen  underwent  postdepositional  crush- 
ing. The  nearly  complete  vertebral  series 
of  the  CM  25362  specimen  has  been  used 
for  the  construction  of  the  remaining  trunk 
and  caudal  region  in  the  model  of  the  A. 
uintaensis  vertebral  column.  The  trunk 
centra  of  CM  25362  have  been  arranged 
according  to  basapophyseal  angles  that  de- 
crease from  180  to  46  degrees,  as  in  A.  calva. 
Decreasing  size  was  used  to  arrange  the 
caudal  vertebrae. 

Although  A.  uintaensis  occurs  much  ear- 


Table  14.     Angle  of  basapophyses,  length,  height,  and  width  of  vertebrae  of  Amia 
uintaensis  compared  with  type  specimens  of  synonymized  taxa  as  illustrated 

IN  figure  22 


Relative 
Vertebral 
Number 


Specimen 


Angle  of 

Basapophyses 

( Degrees ) 


Length 
(mm) 


Height 

(mm) 


Width 

(mm) 


6 


A.  uintaensis  PU  10101 
A.  uintaensis  PU  16236 
P.  uintaensis  ANSP  8044 
P.'sp.  USNM  170973 

A.  uintaensis  PU  10101 
A.  uintaensis  PU  16236 
A.  whiteavesiarm  NMC  6197 
P.  sp.  FMNH  P27465 

A.  uintaensis  PU  10101 
A.  uintaensis  PU  16236 
P.  sp.  USNM  3966 
P.  medius  USNM  3959 

A.  uintaensis  PU  10101 
A.  uintaensis  PU  16236 

A.  uintaensis  PU  10101 
A.  uintaensis  PU  16236 
A.  uintaensis  PU  13865 

A.  uintaensis  PU  10101 
A.  uintaensis  PU  16236 
P.  uintaensis  ANSP  5558 


ISO" 

180° 

180° 
180° 

180° 


8.0 

32.0 

45.0 

6.0 

31.0 

44.0 

8.0 

32.0 

46.0 

5.5 

19.0 

29.0 

8.0 

32.0 

44.5 

8.5 

28.5 

39.0 

8.5 

29.0 

40.0 

9.0 

28.0 

36.0 

10.0 

33.0 

43.0 

8.0 

30.0 

41.5 

8.5 

21.5 

29.0 

8.5 

22.0 

30.0 

10.0 

33.0 

44.0 

7.5 

33.0 

40.8 

11.0 

33.5 

44.0 

9.5 

31.5 

39.0 

4.5 

16.5 

21.5 

11.0 

34.0 

42.5 

9.0 

33.5 

34.0 

10.5 

32.5 

40.0 

Fossil  Amiids  •  Boreske        55 


licr  in  time  than  A.  calva  and  A.  scutata,  it  liodicd  form  tlian  its  contemporary,  A. 
has  approximately  the  same  total  number  of  fra^osa,  which  has  a  mean  of  65  centra, 
centra    (85),    and    like   them    is   a   longer-      Tlu>  vertebral  column  of  A.  umfaenm  does, 


B 


4.  uintaensis 
PU  lOIOI 


A.  uintaensis 
PU  16236 


P.  uintaensis 
ANSP  8044 


A .  whiiteavesiana 
NMC  6197 


R  sp. 
USNM    3966 


A.  uintaensis 
PU  13865 


P.  sp. 
USNM   170973 


P  sp. 
FMNH  P27465 


R  uintaensis 
ANSP 5558 


P  medius 
USNM  3959 


Fig.   22.      First  anterior  trunk  vertebrae   (A,B)  of  Amia   uinfaens'is  compared   with  type  specimens  of  synonymized  taxa 
(refer  to  Table  14  for  data). 


56 


Bulletin  Museum  of  Comparative  Zoologij,  Vol.   146,  No.   1 


11 


R  sp.  P.  sp. 

USNM  170973       USNM  3962 


P.    Sp. 
USNM  170973 


"-^,/ 


P.  plicotus 
AMNH2539 


P  sp. 
PU  20523 


P  Sp. 
USNM  170973 


P  sp. 
USNM  170973 


12C 


12d 


i 


Amia.  sp. 
ANSP2337 


P.  plicafus 
USNM  170974 


P.  medius 
USNM  3959 


12a 


p.  medius  P.   sp. 

YPM  6238       FMNH  P27465 


P.  medius 
USNM  3959 


12b 


Amia  sp. 
ANSP  2339 


Fig.   23.      Seventh    through    fourteenth    mid-trunk    vertebrae    of    Amia    uinfaensis    compared    with    type    specimens    of 
synonymized  taxa  (refer  to  Table  15  for  data). 


Fossil  Amiids  •  Borcske        57 


however,    differ   meristieally   from    that   of  36  (mean)   in  A.  sctitata.    The  number  of 

A.  calva  and  A.  scutata  in  number  of  verte-  diplospondylous  vertebrae  is  20-21,  as  eom- 

brae  in  the  various  regions.    There  are  31  pared   with    14-17   in   A.   calva   and    15   in 

trunk  eentra  in  A.  uintacnsis  (PU   13865),  A.  scutata.  This  variation  from  A.  ra/i;«  and 

as  opposed  to  37  (mean)   in  A.  calva  and  A.  scutata  in  the  organization  of  the  verte- 


15 


/?  plicotus     P.  medius 
USNM  3958    USNM  3959 


R  medius 
YPM  6239 


R    sp. 
USNM  3966 


P  symphysis 
PU  10099 


19a 


19b 


P  medius 
USNM  3959 


R  sp. 
USNM  3966 


R  sp 
USNM  3962 


P  sp.  P  sp. 

USNM  3966        FMNH  P27465 


P  medius 
YPM  6240 


P  sp. 
USNM  3966 


19C 


R  sp.  R  sp- 

FMNH   P27465       USNM  3963 


P.    sp.  P  medius 

USNM  3966      USNM  3959 


R  medius 
USNM  3959 


R   loevis 
USNM  3968 


P  uintaensis 
ANSP  3151 


Fig.    24.      Fifteenth    through    twenty-second    posterior    trunk    vertebrae    of    Amio    uiniaemis    compared    with    type    speci- 
mens of  synonymized  taxa  (refer  to  Table  16  for  data). 


5(S         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


26 


27 


28 


29 


30 


31 


Psp. 
YPM  6242 


/?  uintaensis 
ANSP3I5I 


P.    loevis 
PU  10109 


R  laevis 
PU  10109 


P.  media 
ANSP5632 


P    Sp. 
USNM  3963 


Amia  sp. 
ANSP2338 


P  symphysis 
PUIOIIO 


A.  mocrospondyla 
NMC  6198 


34 


37 


39 


43 


44 


45 


50 


51 


59 


m 


P.  laevis 
USNM  3968 


P   sp. 
FMNH  PF4509 


P    sp. 
USNM  3966 


P.  medius 
USNM  3959 


P.   medius 
USNM    3959 


P  sp. 
USNM  5450 


P.   medius 
USNM  3959 


P.  corsoni 
USNM  3961 


Fig.    25.      Posterior    trunk    and    caudal    vertebrae    of    Am/o    umfoensis    compared    with    type    specimens    of    synonymized 
taxa  (refer  to  Table  17  for  data). 


Fossil  Amiids  •  Boreske        59 


Table  15.     Angle  of  hasapophyses,  length,  height,  and  width  ok  vehtehuae  of  Amia 
iiintacn.sis  compared  with  type  specimens  of  synonymized  taxa  as  illustrated 

IN  figuhe  23 


Relative 
Vertebral 
Number 


Specimen 


Angle  of 

Basapophyscs 

( Def^rees ) 


Length 
(mm ) 


Height 
(mm) 


Width 
( mm ) 


7 
8 
9 

11 

12a 

12b 
12c 
12d 
14 


P.  sp.  USNM  170973 
P.  sp.  USNM  3962 

P.  sp.  USNM  170973 

P.  plicatus  AMNH  2539 
P.  sp.  USNM  170973 

P.  .s7>.  PU  20523 

P.  sp.  USNM  170973 

P.  77ic(lius  YPM  6238 
P.  sp.  FMNH  P27465 
P.  mcdius  USNM  3959 

Aiiiiu  sp.  ANSP  2339 

A7nia  sp.  ANSP  2337 

P.  ])licatus  USNM  170974 

P.  mcdius  USNM  3959 


179° 

8.5 

19.5 

25.5 

7.0 

20.0 

28.0 

178° 

8.5 

19.5" 

26.0 

177° 

6.0" 

18.0^ 

24.0 

8.0 

21.0 

25.5 

174° 

7.5 

25.0 

30.0 

7.5 

22.0' 

25.0 

6.0 

19.0 

24.0 

171° 

11.0 

30.0 

35.0 

9.0 

22.0 

28.0 

167° 

10.0 

29.0" 

40.0 

166° 

10.0 

29.0 

38.5 

163° 

8.0 

24.0 

30.0 

160° 

7.5 

19.0 

23.0 

Est. 


Taulk  16.     Angle  of  hasapophyses,  ijiingth,   hkic;ht,  and  width  of  verteijuae  of  Amu/ 
uintactisis  compared  with  type  specimens  of  synonymized  taxa  as  illustrated 

IN  figure  24 


Relative 
Vertebral 
Number 


15 

17 

18 

19 

19a 
19b 

19c 

20 

21 
22 


Sjiccimen 


P.  plicatus  USNM  3958 
P.  mcdius  IfSNM  3959 
P.  mcdius  YPM  6239 
P.  sp.  USNM  3966 
P.  sijmphysis  PU  10099 

P.  mcdius  USNM  3959 
P.  sp.  USNM  3962 

P.  sp.  USNM  3966 

P.  ,v/;.  USNM  3966 
P.  sp.  FMNII  P27465 

P.  i7icdius  YPM  6240 

P.  sp.  USNM  3966 

P.  sp.  FMNH  P27465 
P.  sp.  USNM  3963 

P.  sp.  USNM  3966 

P.  Tticdius  USNM  3959 

P.  mtY/tt/.v  USNM  3959 

P.  /at't;i.v  USNM  3968 
P.  uiiitacrisis  ANSP  3151 


Anglo  of 

Basapophyses 

( Degrees ) 

Length 
(mm) 

Height 
(mm) 

Width 
( mm ) 

156° 

8.5 
8.5 
8.5 
8.5 
5.5 

22.5 
23.0 
22.0 
21.0 
15.5 

25.0 
27.0 
24.5 
25.5 
20.0 

153° 

9.0 
9.0 

24.0 
23.0 

29.0 
28.0 

149° 

9.0 

21.0 

25.5 

143° 

8.5 
11.0 

23.0 
28.0 

27.8 
33.0 

139° 

7.0 

19.0 

23.0 

138° 

— 

— 

136° 

11.0 
7.0 

25.0 
16.5 

35.0 
20.0 

132° 

9.0 
9.0 

21.0 
22.0 

27.0 
26.0 

122° 

9.5 

22.0 

27.0 

117° 

10.0 
12.0 

26.0 
28.0 

30.0 
29.0 

60 


Bulletin  Museum  of  Coniparative  Zoology,  Vol.   146,  No.   1 


Table  17.     Angle  of  basapophyses,  length,  height,  and  width  of 
uintaensis  compared  with  type  specimens  of  synonymized  taxa 

IN  figure  25 


vertebrae  of  Amia 
as  illustrated 


Relative 
Vertebral 
Number 

Specimen 

Angle  of 

Basapophyses 

( Degrees ) 

Length 

( mm ) 

Height 
( mm ) 

Width 
( mm ) 

24 

P.  sp.  YPM  6242 

102° 

9.0 

17.0 

22.0 

26 

P.  tiintaensis  ANSP  3151 

97° 

10.0 

23.0 

28.0 

27 

P.  laevis  PU  10109 

90° 

13.0 

29.0 

33.0 

28 

P.  laevis  PU  10109 

83° 

14.0 

28.0 

30.0 

29 

P.  media  ANSP  5632 

80° 

8.0 

16.0 

18.0 

30 

P.  sp.  USNM  3963 

P.  symphijsis  PU  10110 

62° 

11.0 
6.0 

14.0 

13.5 

31 

Amia  sp.  ANSP  2338 

A.  macrospondyla  NMC  6198 

46° 

13.0 
12.0 

26.0 
25.0 

23.0 
22.0 

34 

P.  laevis  USNM  3968 

6.5 

22.0 

18.0 

39 

P.  sp.  FMNH  PF  4509 

7.0 

19.0 

18.0 

43 

P.  s/;.  USNM  3966 

7.0 

19.0 

18.0 

44 

P.  medins  USNM  3959 

6.0 

16.0 

18.0 

45 

P.  7ne(/ii«  USNM  3959 

5.5 

17.5 

18.0 

50 

P.  s/;.  USNM  5450 

7.0 

15.0 

13.5 

51 

P.  »!«//»«  USNM  3959 

5.0 

17.0 

11.0 

59 

P.  corsonii  USNM  3961 

4.0 

11.0 

10.0 

bral  column  into  region.s  and  types  of  verte- 
brae appears  to  be  a  useful  taxonomic 
character  of  A.  nintaensi.^. 

The  neural,  aortal,  and  haemal  facets  do 
not  appear  to  vary  much  from  those  of 
A.  calva.  The  first  six  ventral  aortal  facets 
show  basically  the  same  pattern  for  both 
species  (Figs.  11,  22).  The  angle  of  basa- 
pophyses in  A.  uintaensis  differs  from  that 
of  A.  calva  in  two  ways.  The  first  six  verte- 
brae all  have  basapophyseal  angles  of  180 
degrees,  and  it  is  not  until  the  seventh 
vertebra  that  these  angles  gradually  begin 
to  decrease.  Because  of  this  more  posterior 
beginning  in  the  decrease  of  the  angles  and 
because  there  are  fewer  tinmk  vertebrae,  the 
rate  of  decrease  of  the  basapophyseal  angle 
is  greater.  These  angles  range  from  ISO 
degrees  anteriorly  to  approximately  45  de- 
grees posteriorly,  about  the  same  as  the 
range  for  A.  calva. 


The  intracolumnar  variation  in  centrum 
shape  seen  in  the  vertebral  column  of  Re- 
cent A.  calva  also  occurs  in  A.  uintaensis 
( Fig.  14).  In  some  respects  the  latter  shares 
certain  characteristics  with  A.  calva.  The 
first  centrum  is  broad  and  thin,  and  usually 
lacks  basapophyses  (Fig.  22).  However, 
centra  between  the  fourth  and  twentieth 
vertebrae  begin  to  acquire  an  almost  sub- 
triangular  outline,  as  opposed  to  the  sub- 
elliptical  form  of  the  A.  calva  trunk  centra 
(Fig.  12).  The  subtriangular  shape  may  be 
a  function  of  greater  size  of  the  centra. 

The  chordal  foramen  is  open  in  all  known 
vertebrae  of  A.  uintaensis  from  the  Paleo- 
cene.  Eocene,  and  Oligocene,  but  is  often 
filled  with  detritus  during  fossilization. 
Estes  (1964:  42)  observed  that  Cretaceous 
specimens  as  well  as  the  Late  Faleocene 
specimen  (PU  16236)  had  the  chordal 
foramen  smoothly  closed  with  bone.    A  re- 


Fossil  Amiids  •  Boreske 


61 


examination  of  PU  16236  reveals  that  the 
chordal  foramen  is  actually  filled  with  fine 
sediment  rather  than  bone,  so  that  the 
character  of  the  closed  foramen  can  only  be 
applied  to  the  Cretaceous  specimens. 
Chordal  foramen  position  in  all  specimens 
shows  slight  intracolumnar  variation  along 
the  tiimk  as  in  A.  calva,  although  occurring 
more  dorsally.  In  the  caudal  region  there  is 
virtually  no  difference  between  the  two 
forms. 

Leidy  characterized  "Protamia"  uintaensis 
on  the  basis  of  five  centra  and  one  basioc- 
cipital.  His  height/ width  proportions  were 
described  in  relation  to  those  in  an  un- 
diagnostic  intracolumnar  standardization  of 
the  centra  of  the  A.  calva  vertebral  column. 
My  measurements  of  the  anterior  trunk 
centra  reveal  that  the  holotypc  ANSP  5558 
has  a  width  1.3  times  the  height,  and  para- 
type  ANSP  8044  has  a  width  1.6  times  the 
height.  Other  paratype  centra  are  posterior 
trunk  centi-a  with  width/ height  ratios  of 
approximately  1:1.  Romer  and  Fryxell 
(1928:  521)  described  a  displaced  posterior 
trunk  centrum  as  having  a  height  of  10  mm 
and  a  width  of  12.5  mm,  about  the  same  as 
in  ANSP  5558.  Estes  (1964:  43),  in  his 
discussion  of  the  height/ width  proportions 
of  A.  uintaensis  centra,  misinterpreted 
Leidy 's  (1873a,  1873b)  diagnosis  of  "Proto- 
mia'  uintaensis  and  Romer  and  Fryxell's 
(1928)  diagnosis  of  "Paramiatus  ^wleyij' 
indicating  that  vertebrae  of  tlie  former  were 
three  times  as  wide  as  deep,  those  of  the 
latter  two  times.  Estes  was  correct,  how- 
ever, in  his  assumption  that  there  is  intra- 
columnar variation  in  height/ width  ratios. 

The  general  pattern  of  intracolumnar 
variation  in  the  A.  uintaensis  vertebral 
column  is  quite  similar  to  that  of  A.  calva; 
there  is  the  same  trend  from  horizontally 
elliptical  centra  to  circular  or  vertically 
elliptical  centra  ( Fig.  14 ) .  Thus  the  earlier 
diagnoses  of  A.  uintaensis  using  height/ 
width  ratios  that  attributed  the  proportions 
of  the  anteriormost  trunk  vertebrae  to  the 
entire  column  are  undiagnostic. 

On  the  basis  of  isolated  centra  and  skull 
material,  the  most  commonly  used  character 


in  differentiating  A.  uintaensis  from  A.  calva 
has  been  the  former's  greater  size.  How- 
ever, the  articulated  specimen  (PU  13865), 
which  is  the  smallest  known  A.  uintaensis, 
is  only  146  mm  longer  than  A.  fra^osa 
(FMNH  2201)  and  16  mm  longer  than  the 
largest  A.  calva  known  to  me  (UMMZ 
197683).  Estes  (1964)  suggested  that  the 
widening  of  the  A.  uintaensis  vertebrae 
might  be  a  function  of  its  greater  size; 
Gould's  (1966)  statement  that  internal 
elements  generally  increase  at  allometric 
rates  to  provide  sufficient  surface  area  to 
maintain  the  external  surface  area  offers  a 
partial  explanation  as  to  why  the  large  A. 
uintaensis  vertebrae  have  greater  width  in 
proportion  to  height  than  they  do  in  smaller 
amiid  vertebrae. 

Discussion 

Two  species  of  Protamia,  one  species  of 
Ilypamia,  six  species  of  Pappichthys,  and 
three  species  of  Amia  have  been  described 
on  vertebral  characters  from  isolated  centra 
and  disarticulated  cranial  elements  (Table 
19).  With  the  exception  of  Amia  ichiteaves- 
iana,  A.  selwyniana,  and  A.  macrosponclyla 
from  the  Oligoeene  Cypress  Hills  Forma- 
tion of  Alberta,  all  these  taxa  are  based  on 
material  from  the  Bridger  Basin,  Bridger 
Formation,  of  Wyoming.  Each  of  these  12 
taxa  will  be  re-evaluated  in  the  following 
discussion.  Of  the  twelve  species  and  four 
genera,  'Trotamia"  uintaensis  (Leidy, 
1873a)  is  the  oldest  name.  Leidy's  type 
specimens  are  all  trunk  vertebrae.  The 
holotypc  ANSP  5558  (Fig.  22)  is  approxi- 
mately the  sixth  anterior  vertebra  and 
displays  the  characteristic  subtriangular 
outline  of  other  specimens.  The  paratypes 
include  trunk  vertebrae  ( ANSP  8044,  3151 ), 
and  a  large  basioccipital  (ANSP  5622).  The 
holotypc  vertebrae  and  the  basioccipital 
are  considered  diagnostic  for  Amia  uintaen- 
sis, on  the  basis  of  their  possessing  the 
characteristic  subtriangular  vertebral  out- 
line, and  a  kidney-shaped  articular  surface 
of  the  basioccipital. 

Leidy  (1873a)  described  Protamia  media 
from    two    large    trunk    centra    from    the 


62         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Bridger  Formation  of  Wyoming.  His  main 
criterion  for  distinguishing  this  form  from 
A.  calva  and  from  the  other  species  of 
"Protarnia"  was  that  the  vertebrae  were 
twice  the  size  of  A.  calva  vertebrae  and 
"somewhat  smaller  than  Protamia  uintaen- 
sis"  (Leidy,  1873b).  The  holotype  USNM 
2181  appears  to  be  from  the  anterior  trunk 
region  ( approximately  the  seventh  or  eighth 
centrum,  as  suggested  by  its  proportions 
and  configurations  of  aortal  facets).  The 
basapophyseal  angle  is  approximately  178- 
180  degrees.  The  paratype  ANSP  5632  is 
from  the  posterior  trunk  region,  with  an  80- 
degree  basapophyseal  angle,  which  is  ap- 
proximately equivalent  to  the  twenty-ninth 
centrum  in  A.  uintaensis  (Table  17;  Fig. 
25).  Cope  ( 1884,  plate  4,  figs.  7-20)  figured 
"PappiclitJiys  mcdius"  on  the  basis  of  14 
disarticulated  centra  from  the  same  locality 
(USNM  3959).  Eight  of  these  are  from  the 
trunk  region  and  correspond  to  centra 
within  the  anterior  to  mid-trunk  region  of 
A.  uintaensis  (Tables  15-16;  Figs.  23-24). 
The  remaining  six  centra  correspond  to  cen- 
tra in  the  caudal  region  (Tables  16-17; 
Figs.  24-25).  Cope  gave  no  description,  but 
in  figuring  these  specimens  he  allocated  to 
them  his  own  genus,  emending  Leidy's 
(1873a)  prior  nomenclature.  Both  Leidy 
and  Cope  had  apparently  assumed  that  the 
characteristics  of  one  or  a  few  vertebrae 
represented  those  of  the  entire  coliunn. 
Both  species  fall  well  within  the  size  range 
of  A.  uintaensis  (Tables  14-17),  and  are 
here  considered  synonyms  of  the  latter. 

Leidy  (1873a)  describc-d  Uypamia  ele- 
gans  from  one  small  trunk  vertebra.  He 
characterized  this  form  as  possessing  a  cen- 
trum that  was  characteristically  "short  in 
proportion  with  its  breadth,  and  it  presents 
sutural  impressions  for  a  contiguous  pair  of 
neural  arches"  (Leidy,  1873b).  ANSP  5580 
appears  to  be  from  the  mid-trunk  region, 
comparable  to  approximately  the  nineteenth 
centrum  as  suggested  by  its  proportions  and 
configuration  of  aortal  facets.  The  basapo- 
physeal angle  is  138-139  degrees.  These 
character-states  and  the  small  size  are  not 


unique,  occurring  as  the  do  in  all  the  other 
species  of  Amia;  Hypamia  elegans  is  there- 
fore a  nomen  duhium. 

Cope  (1873)  described  Pappiclithys 
plicatus  from  the  anterior  portion  of  a  large 
left  dentary  (AMNH  2539).  Other  type 
material  included  two  premaxillae,  a  right 
quadrate,  a  left  epihyal,  an  anterior  portion 
of  an  ectopterygoid,  three  trunk  vertebrae, 
and  numerous  fragments  of  angulars.  He 
characterized  this  form  primarily  on  the 
basis  of  dermal  sculpture  of  the  "cranial 
fragments  being  roughly  grooved."  The 
angular  in  A.  uintaensis  is  generally 
marked  by  more  pronounced  dermal  sculp- 
tiu-e  than  the  other  mandible  elements. 
His  diagnosis  of  the  vertebrae  (USNM 
3958)  is  based  on  proportions  and  mor- 
phology of  neural  and  aortal  facets,  both 
of  which  correspond  to  various  trunk  ver- 
tebrae in  A.  uintaensis  (Tables  15-16; 
Figs.  23-24).  The  description  of  the  re- 
maining elements  conforms  with  other 
elements  of  A.  uintaensis.  Pappichthys 
plicatus  is  tlierefore  a  synonym  of  the  latter. 

Cope  (1873)  described  Pappichthys 
sclerops  from  a  large  left  dentary.  He 
characterized  this  form  as  possessing  a 
dentary  "more  compressed  and  deeper"  than 
that  in  A.  calva  and  other  species  of  "Pap- 
pichthys." The  dentary  (USNM  3965)  in 
all  respects  greatly  resembles  all  dentaries 
that  have  been  referred  to  A.  uintaensis,  and 
I  regard  Pappichthys  sclerops  as  a  synonym 
of  the  latter. 

('ope  (1873)  described  Pappichthys 
laevis  from  a  large  anterior  dentary  frag- 
ment (USNM  3968).  Other  type  materials 
include  a  premaxillary  fragment,  fragments 
of  angulars  (AMNH  2570),  a  left  quadrate 
fragment,  a  trunk  vertebra  fragment,  and  a 
caudal  vertebra.  Although  Cope  distin- 
guished this  taxon  from  other  species  of 
Pappichthys  on  vertebral  proportions,  vari- 
ances in  dermal  sculpture,  dentary  alveolar 
count,  and  obliqueness  of  alveolar  face, 
these  character-states  occur  in  A.  uintaensis. 
PappicJitJujs  laevis  is  therefore  a  synonym 
of  the  latter. 


» 


Fossil  Amiids  •  Boreske 


63 


Cope  (1873)  described  Pappichthys  sym- 
pJu/sis  from  two  large  fragments  of  trunk- 
vertebrae  and  a  iiral  (USNM  3960).  His 
diagnosis  rests  primarily  on  eonfiguration 
of  neural  faeets  and  basapophyseal  length. 
Osborn  et  cil  ( 1878:  104)  later  reported  two 
eaudal  vertebrae  as  cotypes  (PU  10099, 
10110).  Cope  (1873)  described  Pappich- 
thys  corsonii  from  12  centra  (USNM  5475- 
5476),  a  basioccipital  (USNM  5476),  and 
a  left  dentary  fragment  (USNM  3961).  He 
distinguished  this  form  from  Pappiclithys 
sympliysis  on  different  neural  facet  mor- 
phology, basapophyseal  length,  and  height/ 
width  proportions.  Merrill  (1907:  14)  cites 
^'PappicJitJujs  sympliysis  =  Pappichthys  cor- 
sonii' without  further  discussion.  The  cen- 
tra of  both  forms  conform  to  centra  in  the 
vertebral  column  of  A.  uintaensis  (Table 
17;  Fig.  25)  and  the  characters  assigned  to 
the  dentary  and  basioccipital  of  Pappich- 
thys corsonii  are  also  found  in  A.  uintaensis; 
thus  both  P.  sympliysis  and  P.  corsonii  arc 
synonyms  of  A.  uintaensis. 

From  the  Early  Oligocene  Cypress  Hills 
Formation,  Saskatchewan,  Cope  (1891) 
described  Amia  wliiteavesiana  from  an  an- 
terior vertebra  (NMC  6197),  and  Amia 
macrospondyJa  from  a  caudal  vertebra 
(NMC  6198).  Both  these  forms  were 
founded  on  variations  of  vertebral  charac- 
ters (height/ width  proportions,  lack  of 
basapophyses,  and  chordal  foramen  posi- 
tion) that  are  also  represented  in  the  verte- 
bral column  of  A.  uintaensis.  The  type 
centrum  of  A.  tiJiiteavesiana  corresponds 
approximately  to  the  second  anterior  verte- 
bra in  A.  uintaensis  (Table  14;  Fig.  22), 
that  of  the  type  centrum  of  A.  macrospon- 
(Jyla  with  the  thirty-first  centrimi  in  A. 
uintaensis  (Table  17;  Fig.  25).  Prior  to 
the  appearance  of  Cope's  ( 1891 )  publica- 
tion. Ami  (1891),  in  his  review  of  the 
Cypress  Hill  fauna,  mistakenly  listed  A. 
whiteavesiana  under  the  name  A.  selwyni- 
ana.  A.  macrospondyhi  and  A.  ivhiteavesi- 
ana  are  here  considered  synonyms  of  A. 
uintaensis;  A.  sehcyniana  is  a  iiornen 
nudum. 


Comments  on  European  and 
Asian  Forms 

Janot  ( 1967 )  described  a  large  amiid, 
Amia  rohusta,  from  the  Late  Paleocene  of 
France,  on  the  basis  of  disarticulated 
material.  She  distinguished  this  form  from 
A.  calva  and  A.  russeUi  on  the  angle  of  the 
ventral  border  of  the  dentary  face,  and  on 
morphology  of  the  parasphenoid  tooth- 
bearing  surface  in  addition  to  other  minor 
morphological  differences.  Many  of  the  di- 
agnostic elements  or  associations  on  which 
A.  uintaensis  is  based,  such  as  coronoid  and 
vomerine  teeth,  regional  vertebral  counts 
and  dorsal  cranial  elements,  are  missing  in 
her  material.  The  elements  that  she  does 
figiue,  however,  closely  resemble  the  com- 
parative bones  in  A.  uintaensis.  Simi- 
larities include  rounded  distal  ends  of 
branchiostegal  rays  (also  in  A.  fragosa), 
subtriangular  morphology  of  trunk  verte- 
brae, extensive  surface  of  parasphenoid 
tooth-patch,  and  shallow  orbital  notch  in 
frontal  (also  in  A.  scutata  and  A.  calva). 
These  marked  similarities  suggest  that  A. 
rohusta  is  a  synonym  of  A.  uintaensis. 
Current  work  on  the  relationship  of  the 
North  American  and  European  continents 
in  the  Early  Cenozoic  ( McKenna,  1972) 
indicates  that  they  were  connected  until  tlie 
Early  Eocene  and  that  there  is  great  sim- 
ilarity between  the  Paleocene  and  Early 
Eocene  mammalian  taxa  at  that  time. 
There  is  thus  no  zoogeographic  problem  in- 
herent in  synonymizing  these  two  species. 

Hussakof  (1932)  described  Pappichthys 
mongoliensis  from  disarticulated  elements 
from  the  Late  Eocene  Ulan  Shireh  beds  of 
the  Shara  Murun  region.  Inner  Mongolia 
(collected  by  the  American  Museum  Cen- 
tral Asiatic  Expeditions.)  At  the  time  of 
Hussakof's  description,  this  collection 
(AMNH  6372)  represented  the  most  exten- 
sive material  of  ''Pappichthys."  The  collec- 
tion includes  numerous  dentaries,  maxillae, 
three  gulars,  three  opercula,  three  cleithra, 
an  hyomandibular,  a  supracleithrum,  a 
vomer,   and   trunk   and   caudal   vertebrae. 


64         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Hussakof  distinguished  this  form  from  A. 
calva  by  the  length  of  the  dentaries  and 
the  morphology  of  the  operculum,  and 
from  species  of  "Pappichthys"  and  "Pro- 
tamia"  on  the  basis  of  comparison  of  verte- 
bral size.  A  comparison  of  the  Mongolian 
material  with  A.  uintaensis  shows  some  dis- 
similarities, but  there  is  still  a  closer  affinity 
between  this  form  and  A.  uintaensis  than 
with  the  other  species  of  Amia.  The  vomer 
bears  numerous  sharp  vomerine  teeth;  the 
hyomandibular  is  deeply  arched,  and  the 
lingual  face  of  the  dentaries  conforms  to 
that  of  A.  uintaensis.  The  dentary,  however, 
is  quite  elongated  anteriorly,  the  supra- 
cleithrum  is  narrower,  and  the  dorsal 
border  of  the  operculum  is  short  and 
ascends  at  a  30-degree  angle  rather  than 
being  horizontal  as  in  A.  uintaensis  (and 
in  other  Amia  species).  The  extrascapular 
is  narrow  and  tapered  to  a  point  rather  than 
flattened  medially.  Thus,  although  Pappich- 
thys  mongoliensis  is  similar  to  A.  uintaensis 
in  many  features  and  is  clearly  related  to  it, 
it  also  differs  in  some  respects.  It  un- 
doubtedly belongs  to  the  genus  Amia,  and 
retention  of  all  the  Mongolian  specimens  in 
Amia  mongoliensis  seems  the  most  practical 
alternative  at  this  time.  The  Mongolian 
higher  vertebrate  taxa  indicate  that  the 
Turgai  Straits  at  least  partially  isolated 
Mongolia  from  Europe  during  at  least  part 
of  the  Cretaceous,  Paleocene,  and  Eocene, 
and  that  probably  little  exchange  took  place 
until  the  Late  Eocene  (Szalay  and  Mc- 
Kenna,  1971:  280-281).  It  may  be  possible 
that  A.  mongoliensis  evolved  from  A. 
uintaensis  during  this  migration. 

Amia  cf.  uintaensis 

Hypodigm.  Cretaceous.  Lance  Forma- 
tion, Wyoming:  CM  256,  YPM  6311,  trunk 
vertebrae;  UCMP  56276,  two  fragments  of 
a  single  vertebra;  UCMP  56277,  one  com- 
plete vertebra,  one  vertebral  fragment,  one 
left  maxillary  fragment.  Hell  Creek  Forma- 
tion, Montana:  AMNH  6385,  trunk  vertebra; 
MCZ  9334,  dentary  tooth  tips.  Aguja  For- 
mation, Texas:  UMM  collections,  maxillary 


fragment.     Ojo    Alamo    Formation,    New 
Mexico:  USNM  collections,  trunk  vertebra. 

Discussion 

Cretaceous  specimens  of  large  amiids 
occur  in  both  Lance  and  Hell  Creek  for- 
mations and  consist  mostly  of  isolated 
and  broken  centra,  and  teeth  that  have 
been  identified  primarily  on  the  basis  of 
size.  The  characteristic  subtriangular  out- 
line of  the  trunk  vertebrae  is  even  more  pro- 
nounced in  these  Cretaceous  specimens, 
wherein  the  lateral  centrum  walls  between 
the  basapophyses  and  the  aortal  facets  are 
concave  ( Fig.  26 ) .  The  chordal  foramen  is, 
as  Estes  ( 1964:  42)  noted,  closed  with  bone, 
as  are  one-third  of  the  vertebrae  referred  to 
A.  fragosa  from  the  Lance  Formation.  How- 
ever, Estes  observed  lateral  concavities  be- 
tween the  neural  facets  and  basapophyses 
in  a  large  vertebral  centrum  (AMNH 
6385)  from  the  Hell  Creek  Formation 
(mistakenly  cited  by  him  as  AMNH  6835 
from  the  Oldman  Formation  of  Alberta). 
Estes  apparently  confused  neural  with 
aortal  facets  and  thus  figured  the  vertebra 
upside  down.  Correct  orientation  of  the 
centrum  (Fig.  26)  shows  concavities  be- 
tween the  basapophyses  and  the  aortal 
facets.  Thus,  Estes  was  incorrect  in  con- 
cluding that  A.  fragosa,  A.  calva,  and  the 
Eocene  specimens  of  A.  uintaensis  "also 
seem  to  lack  the  concavity  between  the 
'basapophysis'  and  neural  arch  present  in 
the  large  Cretaceous  specimens."  Two 
other  specimens  from  the  Lance  Formation 
(YPM  6311,  CM  256;  Fig.  26)  also  show 
the  prominent  concavities  between  the 
basapophyses  and  aortal,  rather  than  neu- 
ral, facets.  In  addition  to  the  vertebrae, 
Estes  described  a  maxillary  fragment  as 
being  larger  and  more  robust  than  that  of 
A.  fragosa,  although  "characteristically 
amiid  in  tooth  implantation  and  general 
shape."  A  more  complete  maxillary  frag- 
ment (UMM  collections)  from  the  Aguja 
Formation  ( Big  Bend  National  Park, 
Brewster  County,  Texas)  conforms  with 
Estes'  (1964)  description. 


Fossil  Amiids  •  Borcnke        65 


B 


Hlii 


H 


Fig.  26.  Comparison  of  different  Cretaceous  vertebrae.  Am'ia  cf.  uinfaensh:  A,  anterior  trunk  vertebra,  CM  256, 
Lance  Formation,  Wyoming;  B,  posterior  trunk  vertebra,  AMNH  6385,  Hell  Creek  Formation,  Montana;  C,  mid-trunk 
vertebra,  YPM  6311,  Lance  Formation,  Wyoming.  Chondrichthyes:  D,  E,  G  (thin  section),  trunk  vertebrae,  FHKSCM 
13024-9,  Black  Creek  Formation,  North  Carolina;  F,  trunk  vertebra,  MCZ  12879,  Peedee  Formation,  North  Carolina. 
Cetacean:  H,  caudal  vertebra,  FHKSCM  13025,  Calvert  Formation?,  North  Carolina. 
1  rz  dorsal,  2  ^  articular  surface,  3  ::=  ventral 


66 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Only  three  new  centra  and  a  maxillary 
fragment  have  been  identified  since  Estes' 
(1964)  study.  The  vertebrae,  as  noted 
above,  differ  in  certain  minor  respects  from 
the  Paleocene  and  Eocene  specimens. 
Whether  or  not  this  material  actually  repre- 
sents A.  uintaensis  or  an  earlier  stage  of 
evolution  can  only  be  determined  when 
more  complete  Cretaceous  material  is  avail- 
able. 

Amia  scutata   Cope,    1875 

Amia  dictycephala  Cope,  1875:  3. 
Amia  exilis  Lambe,   1908:    12. 

Holotijpe.  USNM  5374,  incomplete  spec- 
imen lacking  the  head  and  body  anterior  to 
the  middle  of  the  dorsal  fin;  anal  and  part 
of  dorsal  and  caudal  fins  well  preserved. 

Type  locality  arul  horizon.  Florissant, 
Colorado.  East  half  of  section  2,  T  13  S, 
R  71  W,  Teller  County,  Colorado;  Flor- 
risant  Formation. 

Age  Range.  Chadronian  (Early  Oligo- 
cene)  to  Orellan  (Middle  Oligocene). 

Hypodigm.  Oligocene.  Cypress  Hills 
Formation,  Saskatchewan:  NMC  6200,  6205, 
vertebrae;  NMC  6201,  basioccipital.  Chad- 
ron  Formation,  South  Dakota:  PU  17172, 
left  dentary  with  posterior  coronoid  bearing 
teeth,  and  a  trunk  vertebra.  Lower  Brule 
Formation,  South  Dakota  and  Nebraska: 
FMNH  PF4508,  PF4509,  CM  3814,  verte- 
brae; FMNH  PF4506,  right  vomer  bearing 
teeth.  Florissant  Formation,  Colorado:  PU 
10172,  nearly  complete  specimen  (counter- 
part =  YPM  6243,  anterior  half;  USNM 
4087,  caudal  half);  YPM  6241,  complete 
caudal  region  (with  counterpart);  UMMP 
V-57431,  nearly  complete  specimen;  USNM 
3992,  partial  specimen,  lacking  skull  and 
tail;  AMNH  2802,  nearly  complete  skiill; 
AMNH  2670,  partial  specimen,  lacking 
skull  and  caudal  region;  AMNH  2671, 
caudal  region. 

Known  distribution.  South  Dakota,  Ne- 
braska, Colorado,  and  Saskatchewan. 

Revised  diagnosis.  Vertebral  meristics 
similar  to  those  of  A.  calva,  but  head/ 
standard-length  proportion  is  intermediate 
between  that  of  A.  uintaensis  and  A.  calva. 


Extrascapular  thicker  at  distal  end  than  in 
A.  calva,  with  concave  posterior  border. 
Pterotic  more  similar  to  that  of  A.  uintaen- 
sis than  of  A.  calva;  anterior  portion  narrow 
and  extended  laterally  to  the  frontal.  Or- 
bital excavations  more  marked  than  in  A. 
calva,  but  not  as  deep  as  in  A.  uintaensis  or 
A.  fragosa.  Preoperculum  resembles  that  of 
A.  uintaensis  more  than  that  of  A.  calva, 
being  narrower  dorsally  than  ventrally. 
Symphyseal  incurving  of  the  dentary  less 
than  in  A.  calva,  but  greater  than  in  A. 
uintaensis.  Ventroposterior  process  of 
cleithrum  heavily  sculptured  as  in  A.  fragosa 
and  A.  uintaensis.  Infraorbital  4  larger 
than  infraorbital  5  as  in  A.  fragosa  and  A. 
uintaensis.  Ossification  of  cranial  bones  ex- 
tensive as  in  other  fossil  species,  greater 
than  in  A.  calva.  Greatest  known  standard- 
length  390  mm. 

Introduction 

Cope's  (1875:  3)  description  of  Amia 
scutata  is  based  on  a  specimen  lacking  the 
head  and  body  anterior  to  the  middle  of 
the  dorsal  fin,  from  the  Middle  Oligocene 
Florissant  Fomiation  near  Florissant, 
Colorado.  He  distinguished  this  form 
from  Amia  dictyocephala  (found  in  the 
same  deposit;  Cope,  1875)  and  Amia  calva 
by  its  larger  scales  "of  which  only  seven 
and  a  half  longitudinal  rows  are  visible 
above  the  vertebral  column."  Cope  de- 
scribed A.  dictyocephala  from  two  partially 
complete  specimens  lacking  skidls  and 
caudal  fins  (USNM  3992,  AMNH  2670), 
two  complete  caudal  regions  (AMNH  2671, 
USNM  4087),  and  a  nearly  complete  skull 
(AMNH  2802);  Osborn  et  al.  (1878)  later 
described  another  specimen  of  A.  scutata 
from  the  same  deposit.  Tliis  specimen 
was  more  complete,  consisting  of  an  axial 
skeleton  and  a  crushed  skull.  They  believed 
A.  scutata  to  be  a  valid  form,  differing  from 
A.  calva  in  having  a  proportionately  larger 
head. 

Comparison  of  known  specimens  of  A. 
scutata  revealed  that  the  counterparts  to 
the  specimen  described  by  Osborn  et  al. 
( PU  10172 )  were  separated  and  sold  to  two 


Fossil  Amiids  •  Borcskr        67 


Fig.   27.      A,    Amia    scutata    UMMP    V-57431;    B,    A.    scufafa    PU    10172;    C,    A.    "dictyocephala"    USNM    3392;    D,    A. 
"dicfyocepbala"  AMNH  2670. 


different  miiseiims.  The  caudal  portion  of  and  is  one  of  the  paratypes  used  by  Cope 
the  counterpart  was  found  in  the  National  (1875)  in  his  description  of  A.  dictijo- 
Museum  of  Natural  History  (USNM  4087)      cephala.    The   anterior  region   was   found 


68 


Bulletin  Museum  of  Comparative  Zoologij,  Vol.   146,  No.   1 


unlabeled   at   the   Yale    Peabody    Museum 
(YPM  6243;  Plate  4).i 

In  1967  another  nearly  eompk^te  speci- 
men was  discovered  from  the  same  deposit 
(Fig.  27A)  and  Cavender  (1970:  42)  re- 
ported the  specimen  A.  dictyocepliala  as 
differing  from  A.  calva  in  having  a  larger 
infraorbital  4,  in  the  sculptin-e  of  cleithrum, 
and  "by  its  proportionately  larger  head  and 
orbit,  and  somewhat  shorter  body." 

Fossil  Record 

Other  than  the  Florissant  Formation,  the 
only  deposits  from  which  elements  of  A. 
scutata  can  be  identified  are  the  Cypress 
Hills  Formation  of  Saskatchewan,  Chadron 
Formation  of  South  Dakota,  and  the  Lower 
Brule  Formation  of  South  Dakota  and 
Nebraska.  Becker  (1961:  38)  reported 
amiid  scales  (UMMP  collections)  from 
the  Late  Oligocene  Passamari  Formation 
and  Middle  Oligocene  Grant  Horse  Prairie 
Shale  of  Montana  (Becker,  1962).  Since 
no  specific  characters  for  scales  of  Amia 
have  yet  been  determined,  it  is  best  to 
allocate  this  material  to  Amia  sp.  Skinner 
et  al.  (1968:  415)  has  reported  Amia  .sp. 
vertebrae  (F:AM  42947)  from  the  Early 
Miocene  Turtle  Butte  Formation  of  South 
Dakota.  Only  two  specimens  were  found; 
since  the  vertebrae  of  A.  scutata  and  A. 
calva  are  morphologically  and  meristieally 
similar.  Skinner  et  al.'s  identification  is  the 
only  possible  one  at  this  time.  The  strati- 
graphic  range  of  A.  scutata  is  therefore  lim- 
ited to  the  Early  and  Middle  Oligocene. 

Description 

Neurocranium..  The  basioccipital  (PU 
10172,  NMC  6201 )  is  similar  to  that  of  A. 
calva.  The  only  available  parasphenoid 
(PU  10172)  is  poorly  preserved,  but  closely 
resembles  that  of  A.  calva  in  length  and 
position  of  ascending  processes. 

The  extrascapular  in  A.  scutata  differs 
slightly  from  that  of  A.  calva  in  that  the 
distal    end    is    relatively    thicker    and    the 


1  The  counterparts  (  USNM  4087,  YPM  6243)  to 
PU  10172  have  been  .subse(iuently  accjuired  by  the 
Museum  of  Natural  History,  Princeton  University. 


L<  orbital  length 
D-  orbital  depth 
^a^dermosphenotlc  angle 


A.  fragosa      D/L=O.I76mn. 


A.  uintaensis    D/L=O.I55mn. 

18° 


Z^"!   40  =  I34<' 


A  scutata      D/L=O.I32mn. 

I5« 


A  calva      D/L"O.IOOmn. 

15° 


IB" 


40=145° 


Z4»r    4.0=135° 


A.   cf.     scutata       O/L'0.121 


[■0=137° 


Fig.   28.      Orbital    dimensions    of    ^m\a    spp. 

posterior  lappet  is  less  pronounced;  also, 
the  posterior  border  is  more  convex  (Fig. 
15).  As  in  A.  calva,  however,  the  proximal 
anterior  corner  is  squared  off,  and  the 
medial  suture  is  relatively  long.  The 
pterotic  in  Ax.  scutata  resembles  that  of  A. 
uintaensis  more  than  that  of  A.  calva  in 
general  morphology,  since  the  anterior  half 
is  narrower  than  the  posterior  half;  in  the 
Recent  species  the  ends  are  nearly  sym- 
metrical. The  anterior  border  extends  fur- 
ther laterally  than  in  A.  calva,  and,  as  in 
A.  uintaensis,  adjoins  the  distal  lateral  side 
of  the  frontal,  rather  than  the  posterior 
border  as  in  A.  calva.  The  dermosphenotic, 
parietal,  frontal,  and  nasal  of  A.  .scutata 
conform  to  these  bones  in  A.  calva.  The 
parietal/ frontal  ratio  is  marginally  within 
the  lower  limit  of  the  range  of  A.  calva 
(Table  7).    The  orbital  excavation  in  the 


Imxssil  Amuus  •   liorcske 


69 


frontal  (Fit:;.  2cS)  is  greater  than  in  A.  caha 
bnt  less  than  in  A.  jruii^osa  or  A.  uintaensis. 
Snprascapulars,  antorbitals,  and  rostrals  are 
not  preserved. 

The  laerinial  is  similar  to  that  of  A.  calva, 
b(>arint!;  a  posterior  noteh  for  the  reeeption 
of  infraorbital  2,  but  in  A.  scutata  the  laeri- 
nial is  more  robust.  Infraorbital  2  and  infra- 
orbital 3  are  similar  to  these  bones  in  A. 
calva.  Infraorbital  4  is  more  massive  pos- 
teriorly than  in  A.  calva;  it  exeeeds  infra- 
orbital 5  in  dorsoventral  length,  and  the 
posterodorsal  corner,  which  in  A.  calva  is 
markedly  acute  is,  in  A.  .scutata,  more 
squared  off.  This  bone  more  closely  re- 
sembles that  of  A.  jraii^osa:  it  is  not  avail- 
ablc>  for  comparison  in  A.  uintacnsis.  Infra- 
orbital 5  is  less  massive  posteriorly  than  in 
A.  calva;  in  this  feature  it  resembles  that  of 
A.  fra'^osa.  It  is  also,  as  in  A.  fra^osa  and  A. 
uintacnsis,  dec>per  anteriorly  than  in  A. 
calva. 

Branchiocranium.  The  supramaxilla  in 
A.  scutata  is  elongated  and  tapered  to  a 
point  anteriorly,  with  a  relatively  straight 
ventral  border  as  in  A.  calva.  It  is  slightly 
longer  and  more  robust  posteriorly,  the 
posterodorsal  border  being  higher  and  less 
obliquely  curved  than  in  A.  calva,  A.  uin- 
tacnsis, and  A.  jra<^osa.  The  preniaxilla 
resembles  that  of  A.  calva.  The  maxilla  is 
wider  posteriorly  and  more  ossified  anteri- 
orly than  that  of  A.  calva,  but  otherwise 
agrees  with  the  bone  in  the  Recent  .species. 

Dermopalatine,  autopalatine,  entoptery- 
goid,  ectopterygoid,  metapterygoid,  and 
vomer  are  not  pr(\served.  Ho\v(^ver,  conical 
vomerine  teeth  are  displayed  on  PU  10172, 
and  resemble  those  of  A.  calva  rather  than 
those  of  A.  fra<s,osa  or  A.  uintacnsis.  The 
relative  munber  of  teeth  and  their  extent 
on  the  vomer  cannot  be  discerned. 

As  in  A.  calva  and  A.  uintacnsis,  the  den- 
tary  in  A.  scutata  lacks  the  dorsal  shelf 
adjacent  to  the  lingual  border  of  the  alveo- 
lar ridge  seen  in  A.  fra^osa  (Fig.  18).  The 
bone  is  very  thick,  especially  toward  the 
mid-lingual  surface,  where  the  dorsal  and 
ventral  halves  meet  to  form  the  Meckelian 
groove.    As  in  A.  uintacnsis  the  upper  wall 


of  this  groove  is  primarily  formed  bv  the 
thickness  of  bone  in  dorsal  half  of  the  lin- 
gual surface;  the  ventral  half  is  barely 
overlain  by  the  dorsal  half.  The  first  coro- 
noid  does  not  extend  past  the  Meckelian 
groove,  and  bears  sharp  conical  teeth,  as  in 
A.  calva  (Fig.  18).  The  anterior  half  of  the 
dentary  is  more  incurved  than  in  A.  uintacn- 
sis, but  not  to  the  extent  that  it  is  in  A. 
calva.  The  anterior  width  of  the  dentary 
also  resembles  that  of  A.  calva  and  A.  uin- 
tacnsis in  that  it  is  evenly  tapered  almost  to 
the  symphyseal  edge.  The  angular  and  sur- 
angular  are  similar  to  comparable  bones  in 
A.  calva,  except  that  they,  like  the  dentary, 
are  more  extensively  ossified.  The  mandi- 
ble/head-length ratio  in  A.  scutata  is  well 
within  this  ratio  range  for  A.  calva  (Table 
7).    The  prearticular  is  not  prc\served. 

The  preoperculuin  is  similar  to  that  of 
A.  uintacnsis,  being  narrower  dorsally  dian 
ventrally,  radier  than  having  both  halves 
rc^latively  equal  in  widdi,  as  in  A.  calva. 
The  operculum  resembles  that  of  A.  calva 
in  morphology  and  opercuhun-depth/oper- 
culum-length  (Table  7).  The  suboperculum 
and  interoperculum  resemble  those  of  A. 
calva  in  general  morphology,  but  the  suture 
between  them  is  longer  anteroposteriorly. 
The  branchiostegal  rays  are  squared  off 
distally,  as  in  A.  calva. 

Post-cranial  skeleton.  The  supracleithnmi 
and  metacleithriun  are  not  preserved.  The 
only  part  of  the  cleithrum  available  for 
study  is  the  ventroposterior  process  in 
UMMP  V-57431  which  in  A.  calva  is  the 
only  area  of  this  bone  that  is  visible  ex- 
ternally. This  region  of  the  cleithrum  in 
A.  scutata  is  heavily  sculptured  (Fig.  21), 
and  as  in  A.  uintacnsis  and  A.  fra^osa,  this 
d(Tmal  ornauKMitation  extends  to  the  edge 
of  the  bone.  In  A.  calva,  diis  d(>rmal  struc- 
ture is  limited  to  the  cent(>r  and  dorsal  re- 
gion of  this  part  of  the  cleithrum. 

Th(>  vertebral  column  of  A.  scutata  re- 
sembles that  of  A.  calva  both  in  number  of 
ccMitra  (Table  9)  and  in  general  morphol- 
ogy of  the  centra.  The  head /standard- 
length  proportion  (0.312)  is  greater  than  in 
A.  calva  (0.271 ),  but  less  than  in  A.  uintaen- 


70         Bulletin  Museuui  of  Comparative  Zoology,  Vol.    146,  No.   1 


sis  (0.322).  The  insertion  of  peetoral  fin/ 
standard-length  and  insertion  of  anal  fin/ 
standard-length  ratios  nrv  both  within  the 
ranges  of  A.  calva,  althongh  the  latter  pro- 
portion for  A.  scutata  is  somewhat  greater 
than  the  mean  for  A.  calva  (  F'ig.  31 ). 

Discussion 

In  the  same  paper  as  his  deseription  of 
Aryiia  scutata.  Cope  (1875:  3)  described 
Amia  clictyocepliala,  also  from  the  Florissant 
Formation.  A.  dictyocephala  was  distin- 
guished from  A.  scutata  by  having  10  to  12 
supravertebral  scale  rows,  and  35  vertebrae 
between  the  anterior  dorsal  fin  pterygio- 
phore  and  the  posterior  anal  fin  ptervgio- 
phore  (USNM  3992  AMNH  2670). "lie 
further  characterized  this  form  from  a  skull 
(AMNH  2S02)  that  "possesses  twelve 
branchiostegal  rays,  and  a  relatively  smaller 
orbit  than  in  Amia  calva."  A  re-examination 
of  these  specimens  in  the  previous  section 
on  meristics  showed  that  Cope's  supra- 
vertebral  scale  row  count  was  in  error,  and 
there  is  no  perceptible  difference  in  this 
feature  between  Recent  and  fossil  Amia 
species  (Table  8).  In  A.  calva,  the  range 
for  the  number  of  centra  between  the  in- 
sertion of  the  dorsal  fin  and  the  terminus 
of  the  base  of  the  anal  fin  is  33-37.  In  the 
type  specimen  of  A.  dictyocephala  (USNM 
3992)  the  number  of  centra  is  35,  and  the 
mean  number  in  specimens  of  A.  scutata  is 
36;  there  is  clearly  no  way  that  this  fcnitiue 
can  be  used  to  distinguish  A.  dictyocephala 
from  A.  scutata  and  A.  calva.  Cope,  on  the 
basis  of  AMNH  2802,  thought  that  an  orbit 
in  A.  dictyocepJiala  was  smaller  than  one  in 
A.  calva,  but  the  small  size  was  due  largely 
to  the  constriction  of  the  orbit  that  resulted 
from  crushing  of  the  dcrmosphenotic  and 
upward  displacement  of  infraorbital  5.  The 
characters  that  Cope  used  to  differentiate 
A.  dictyocephaki  from  A.  scutata  are  un- 
diagnostie,  and  my  studies  of  the  specimens 
show  no  morphological  or  meristie  differ- 
ence; A.  dictyocepJiala  is  here  considered 
to  be  a  synonym  of  A.  scutata. 

Lambe  (1908:  12-13)  described  Amia 
exilis   from    a    single    basioccipital    (NMC 


6201 )  and  two  mid-trunk  \  c>rtebrae  (  NMC 
6200,  6205)  from  the  Farly  Oligocene 
Cypress  Hills  Formation  of  Saskatchewan. 
The  temporal  occurrence  of  these  elements 
is  equivalent  to  that  of  A.  scutata.  Lambe's 
description  of  the  basioccipital  conforms  to 
that  of  A.  scutata  in  being  more  extensively 
o.ssified  than  in  A.  calva.  His  diagnosis  of 
the  two  centra  is  founded  on  height/ width 
proportions,  ehordal  foramen  position,  basa- 
pophyseal  angle,  and  configuration  of  nc>ural 
facets.  Because  A.  scutata  resembles  A. 
calva  in  vertebral  morphology,  the  charac- 
ters that  Lambe  uses  to  distinguish  A.  exilis 
are  undiagnostic;  I  therefore  consider  A. 
exilis  iis  a  synonym  of  A.  scutata. 

Amia  cf.  scutata 

Hypodi^m.  Miocene.  Pawnee  Creek 
Formation,  Colorado:  UCMP  38222,  nearly 
complete  cranial  roof,  infraorbitals  4  and  5, 
nearly  complete  anterior  portion  of  palate, 
two  branchiostegal  rays,  maxillae,  and  right 
dentary. 

Description 

The  general  morphology  of  the  cranial 
roof  resembles  both  A.  scutata  and  A.  calva 
in  parietal/ frontal  ratio  (Table  7),  rectan- 
gular parietals,  and  shape  of  dermosphen- 
otic  and  nasal  (Fig.  29).  The  extrascapular 
more  closely  resembles  that  of  A.  scutata  in 
its  greater  width  and  less  pronoimced  distal 
posterior  lappets.  The  pterotic  also  resem- 
bles that  in  A.  scutata  in  its  being  narrower 
anteriorly  than  posteriorly,  and  in  bordering 
die  frontal  laterally  rather  than  posteriorly. 
The  size  and  depth  of  the  orbital  excavation 
is  intennediate  between  that  of  A.  scutata 
and  A.  calva  (Fig.  28).  The  maxilla  is 
similar  to  that  of  A.  calva,  being  less  robust 
posteriorly  than  that  of  A.  scutata.  The 
branchiostegal  rays  are  squared  off  distally, 
as  are  tliose  of  both  A.  calva  and  A.  scutata. 
Infraorbital  4,  although  posteroventrally  in- 
complete, is  clearly  closer  to  that  of  A.  scu- 
tata than  A.  calva  in  being  relatively  larger 
than  infraorbital  5,  and  in  the  posterodorsal 
corner  being  squared  off  rather  than  acute 
as  in  A.  calva.   Infraorbital  5  resembles  that 


I 


Fossil  Amiids  •  Boreske        71 


of  A.  scutata  in  size  relative  to  infraorbital  4, 
the  anterior  end  bcnng  narrower  than  in 
A.  scutata;  this  featnre  eontributes  to  lessen- 
ing the  relative  width  of  the  orbit.  The 
dentary  resembles  that  of  A.  scutata  in 
being  wider  anteriorly  than  in  A.  calva;  the 
dorsal  lingual  surfaee  only  slightly  overlaps 
the  ventral  lingual  siuface  as  in  A.  scutata 
(Fig.  18);  Meckel's  groove  is  thus  similar 
to  that  of  A.  scutata.  There  is  no  available 
palate  in  A.  scutata  for  comparison.  The 
number  of  vomerine  teeth  is  18  and  21, 
which  is  bracketed  by  the  range  for  A. 
calva  (Estes  and  Berberian,  1969:  5).  As 
Estes  ( 1964 )  noted  for  this  specimen,  these 
teetli  are  sharper  and  more  incurved  ex- 
ternally   than    internally;    this    disparity   is 


more  distinct  in  this  form  than  in  the  extant 
species.  The  hyomandibular,  entopterygoid, 
ectopterygoid,  dermopalatine,  and  pre- 
maxilla  are  poorly  preserved,  but  appear 
to  resemble  these  bones  in  A.  calva. 

Discussion 

Estes  (1964:  36)  and  Estes  and  Tihen 
(1964:  454)  referred  to  this  specimen  as 
Amia  sp.  (and  in  error  gave  the  source  as 
White  River  Formation).  The  .specimen 
resembles  A.  scutata  in  some  elements,  A. 
calva  in  others,  and  is  intermediate  in 
several  character-states,  notably  bone  thick- 
ness and  size  of  orbits.  It  does,  however, 
appear  to  show  a  stronger  resemblance  to 
A.  scutata  than  to  A.  calva,  particularly  in 


Fig.   29.      Amia  cf.  scutata  DC  38222,  Late  Miocene,  Pawnee  Creek  Formation,  Colorado. 


72         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


the  morphology  of  the  extrascapiilar,  ptcro- 
tic,  dentary,  and  infraorbitals  4  and  5,  and 
I  have  thus  compared  it  with  the  fossil 
species.  Since  this  is  a  form  that  is  both 
morphologically  and  temporally  interme- 
diate between  A.  scutata  and  A.  calva,  it  is 
difficult  to  determine  whether  or  not  this 
specimen  actually  represents  A.  scutata  or 
a  later  stage  of  evolution  leading  to  A.  calva, 
but  it  is  at  least  of  interest  in  documenting 
the  slow  phyletic  development  toward  A. 
calva  in  mid-Cenozoic  time. 

Amia  cf.  calva 

Hijpodi^m.  Pliocene.  Lower  Valentine 
Formation,  Nebraska:  UCMP  65851,  an- 
terior portion  of  left  dentary  and  a  trunk 
vertebra;  UMMP  521S7,  right  nasal,  ectop- 
terygoid  fragment,  unidentified  cranial 
fragments;  UMMP  421S5,  right  dentary 
fragment.  Ogallala  Formation,  Kansas: 
UMMP  55574-55578,  three  right  and  two 
left  dentary  fragments;  UMMP  55579,  in- 
complete right  cleithrum;  UMMP  55583,  a 
right  extrascapular;  UMMP  55580,  a  right 
maxilla;  UMMP  55585,  a  left  premaxilla; 
UMMP  55586,  several  scales. 

Discussion 

Smith  (1962),  and  Estes  and  Tihen 
(1964)  described  as  Amia  sp.  a  nasal  and 
dentary,  and  cranial  fragments  from  the 
Lower  Valentine  Formation,  Nebraska. 
Wilson  (1968)  described  as  Amia  calva 
denatry  fragments,  a  premaxilla,  a  maxilla, 
an  extrascapular,  an  incomplete  cleithrum, 
and  several  scales  from  the  Ogallala  Forma- 
tion, Kansas.  This  Early  Pliocene  material 
resembles  A.  calva  more  closely  than  does 
the  Miocene  A.  cf.  scutata  specimen  noted 
above;  the  elements  are  very  lightly  ossified 
as  in  the  Recent  species.  The  cleithrum  is 
distinctly  A.  calva-\ike  in  its  lack  of  distal 
marginal  dermal  sculpture.  The  dentary 
fragments  are  also  thinly  ossified  as  in  A. 
calva,  but  are  slightly  wider  relative  to  the 
dentary  in  the  Recent  species,  as  in  the 
Miocene  form.  Temporally,  this  Pliocene 
material  is  later  than  the  Miocene  form  and 
earlier  than  A.  calva;  morphologically,  how- 


ever, the  available  elements  conform  with 
A.  calva. 

Amiidae  incertae  sedis 

Hypodi^m.  Cretaceous.  Paluxy  Forma- 
tion, Texas:  SMUSMP  62270,  dentary  frag- 
ments, premaxillary  fragment,  vertebrae, 
maxillary  fragments,  and  an  unidentified 
palatal  bone  bearing  teeth;  FMNH  7050, 
basioecipital;  FMNH  7051,  mid-trunk  ver- 
tebra; FMNH  7052,  anterior  trunk  vertebra 
fragment;  FMNH  7053-7054,  anterior  trunk 
vertebrae;  FMNH  7055,  caudal  vertebra; 
FMNH  7056,  small  vertebrae;  FMNH  7049, 
unidentified  palatal  bone  bearing  teeth. 

Description 

The  dentaries  are  fragmentary  (Fig.  30); 
the  only  diagnostic  features  available  for 
comparison  with  other  amiid  forms  are 
related  to  the  anterior  region  of  the  dentary. 
The  surfaces  pits  on  the  exterior  side  of  the 
dentary  are  relatively  larger  and  deeper 
than  in  any  species  of  Amia.  The  dentaries 
lack  the  dorsal  shelf  adjacent  to  the  lingual 
side  of  the  alveolar  ridge  seen  in  A.  fragosa. 
The  coronoid  articulation  surface  descends 
directly  from  the  alveolar  ridge,  as  in  a  Uro- 
cles  dentary  from  the  Late  Jurassic  (Pur- 
beck)  of  England  (BMNH  48236).  The 
lingual  surface  above  the  Meckelian  groove 
is  relatively  short,  even  more  so  than  in 
Amia  uintaensis,  and  the  groove  itself  is 
quite  wide,  more  so  than  in  BMNH  48236. 
The  anterior  portions  of  the  dentaries  are 
relatively  straight,  rather  than  incurved  as  in 
Amia  fragosa,  and  are  evenly  tapered  to  the 
symphyseal  edge.  The  dentary  and  pre- 
maxilla teeth  are  broken,  but  in  dorsal  view 
the  interior  surfaces  of  the  broken  teeth  are 
very  even,  lacking  the  serrated  outline  seen 
in  other  species  of  Amia.  Only  the  anterior 
portion  of  the  premaxilla  is  present;  it  bears 
nine  alveoli,  conforming  in  this  respect  with 
all  Am,ia  species.  The  premaxilla,  although 
incomplete,  displays  the  anterior  (ventral) 
edge  of  the  large  foramen  that  is  character- 
istic of  Amia.  Only  part  of  the  anterior 
maxilla  is  present  in  the  specimens  avail- 
able, and  since  the  more  diagnostic  aspects 


Fossil  A  muds  •  Boreske        73 


occur  posteriorly,  it  is  difficult  to  determine  smaller  fragment  (SMUSMP  62270)  bears 

any   affinities   witli    particular   specic\s;    the  pillar-shaped  teeth  with  nipple-like  tips,  as 

anterior  portions  that  are  available  gener-  in  the  tooth-bearing  palatal  bones  in  species 

ally    conform    with    those    of    Amia.     The  of  Amia.    Posterior   to   the   spinal   arterial 

specific   bones   to   which   the   palatal    frag-  foramina    the    basioccipital    includes    one 

ments    belong    cannot   be    identified.     The  fused   vertebra.    As    in   Amia   fra^osa   and 


Fig.    30.      Amiidae    incerfae    sedis,    Early    Cretaceous,    Poluxy    Formation,    Texas:    A'-A-,    anterior    portion    of    left    den- 
tary;   B,  premaxlllary  fragment;   C,  anterior   portion    of   rigfit  maxilla;    D,    unidentified    palatal   fragment.    XO-15 


74         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Amia  caJva,  the  basioccipital  has  an  ovoid 
articular  surface  with  no  dorsal  indentations 
between  the  neural  facets.  The  large  verte- 
brae are  thickly  ossified,  as  in  the  Creta- 
ceous specimens  of  Amia  cf.  uintaensis. 
The  chordal  foramina  are  closed  and  the 
only  available  large  mid-trunk  centrum  dis- 
plays the  pronounced  triangular  outline 
characteristic  of  Amia  uintaensis.  None  of 
the  large  vertebrae  display  the  character- 
istic Amia  aortal  facets;  they  do,  however, 
possess  neural  facets,  and  the  mid-trunk 
centra  bear  basapophyses.  The  small  verte- 
brae are  also  thickly  ossified  and  the 
chordal  foramina  of  the  trunk  vertebrae  are 
closed.  As  Traquair  (1911:  39)  noted  for 
Amiopsis  cloUoi,  the  lateral  sides  of  the 
vertebrae  are  marked  by  a  number  of  vari- 
able excavations,  or  "oval  fossae"  (Fig.  7). 
These  smaller  mid-trunk  vertebrae,  unlike 
the  large  ones,  display  both  aortal  and 
neural  facets,  as  well  as  basapophyses  and 
lateral  oval  fossae. 

Discussion 

Thurmond  ( 1969 :  88 )  reported  "various 
fragments  of  an  undetermined  amiid"  from 
the  Paluxy  Formation  of  Texas,  which  is 
the  earliest  known  occurrence  of  amiids  in 
North  America.  He  further  noted  that 
amiid  material  occurred  both  in  freshwater 
and  marine  zones  and  that  a  further  descrip- 
tion of  this  material  would  be  the  subject 
of  a  later  study.  He  was  uncertain  as 
to  whether  the  amiids  occurring  in  the 
marine  zones  were  actually  marine  or  were 
freshwater  forms  secondarily  deposited  in 
the  marine  areas.  None  of  the  material  can 
be  referred  to  Atnia  since  it  displays  charac- 
teristics of  Amia  uintaensis,  Amia  fragosa, 
Urocles,  and  Amiopsis,  as  noted  in  the 
above  description.  The  vertebrae  suggest 
the  possibility  of  more  than  one  form:  the 
large  vertebrae  are  subtriangular  and  re- 
semble Am'a  uintaensis  in  morphology, 
with  the  exception  of  the  lack  of  aortal 
facets  on  the  trunk  vertel:)rae.  The  small 
vertebrae  are  Amia  frag,osa-\ike  in  morphol- 
ogy; they  possess  aortal  facets,  but  also 
display  lateral  oval  fossae  characteristic  of 


Amiopsis.  In  reviewing  the  European  Juras- 
sic and  Cretaceous  Urocles,  Lange  (1968) 
found  little  morphological  justification  to 
warrant  continued  generic  distinction  be- 
tween Urocles  and  species  described  by 
Woodward  ( 1916 )  as  belonging  to  Amiop- 
sis from  the  Purbeck  Beds  near  Wevmouth, 
Dorset.  Lack  of  knowledge  of  the  skull  of 
Amiopsis  makes  it  impossible  to  compare 
cranial  elements  with  those  of  other  amiids; 
the  singular  postcranial  feature  charac- 
terizing Amiopsis  is  the  lateral  o\'al  fossae 
of  the  v^ertebrae.  Although  Lange  suggests 
that  both  Amiopsis  and  Ainia  evolved  inde- 
pendently from  different  Urocles  species- 
groups,  it  is  premature  to  attempt  to  do 
more  than  indicate  morphological  similar- 
ities or  dissimilarities  since  the  phylogenetic 
relationship  of  Amiopsis  with  Urocles  or 
Amia  cannot  be  clearly  defined  until  a 
much-needed  review  of  the  taxon  has  been 
completed,  and  until  more  Amiopsis  mate- 
rial is  made  available  for  study. 

The  Paluxy  material  shows  resemblances 
to  two  early  Aryiia  species,  Amia  uintaensis 
and  Amia  fragosa,  as  well  as  to  the  Late 
Mesozoic  European  amiids,  Urocles  and 
Amiopsis.  Whether  the  Paluxy  material 
represents  one  or  more  forms  intermediate 
between  Atnia  and  Urocles  (or  Amiopsis) 
or  whether  it  belongs  to  some  other  group 
of  amiids  that  became  extinct  before  the 
end  of  the  Cretaceous  cannot  be  deter- 
mined, since  taxonomic  evaluation  of  this 
material  is  limited  by  the  lack  of  articulated 
specimens. 

SPECIMENS  REMOVED  FROM 
THE  AMIIDAE 

Miller  (1968:  468-470,  pi.  1,  figs.  1,  3, 
7-9)  questionably  identified  as  Protamia 
sp.  one  large  (FHKSCM  13025)  and  three 
small  centra  (FHKSCM  13024-9)  recovered 
from  a  channel  sandstone  cut  into  the  Up- 
per Cretaceous  Black  Creek  Formation, 
Phoebus  Landing,  North  Carolina.  Since 
all  known  Ainia  are  freshwater  forms  and 
since  these  centra  were  associated  with  vari- 
ous marine  vertebrates.  Miller  (1968:  467) 
concluded  that  the  channel  sandstone  con- 


Fossil  Amiids  •  Boreske 


75 


tained  a  mixed  fauna,  "the  ehannel  sand- 
stone formed  in  an  estuarine  or  tidal  en- 
vironment." 

My  studies  indicate  that  these  specimens 
are  not  amiid.  The  smaller  vertebrae  are 
horizontally  ovoid.  A.  uintaensis  trunk  cen- 
tra have  concavities  between  the  basa- 
pophyses  and  aortal  facets  ( Fig.  26,  A-C ) . 
A  thin  section  (Fig.  26,  G)  through  the 
articular  surface  of  one  of  the  Nortii  Caro- 
lina specimens  (FHKSCM  13924-9)  has  a 
radial  structure  resembling  that  of  S(juatimi 
and  other  sharks  (Hasse,  1882,  tables  17- 
18).  All  layers  are  laminated  parallel  to  the 
exterior  surface  and  are  crossed  by  various 
perpendicular  vascular  foramina.  Their 
articular  surfaces  are  slightly  concave,  while 
those  of  Ainia  are  markedly  so.  Each  of  the 
small  vertebrae  bear  horizontal  basapoph- 
yses  as  in  Recent  Sqiialus,  and  are  best 
referred  to  the  elasmobranchs. 

The  large  vertebra  is  a  cetacean  caudal 
(Fig.  26,  H),  possibly  belonging  to  the 
Cetotheriidae  (Clayton  Ray,  1971,  personal 
communication).  The  centrum  is  ovoid, 
with  very  slightly  concave  articular  surfaces, 
and  lacks  a  chordal  foramen,  as  well  as 
ventral  facets.  The  dorsal  facets  for  the 
accommodation  of  metapophyses  are  well 
defined.  Since  this  centrum  is  from  a  ma- 
rine mammal,  it  is  more  probably  from  the 
Miocene  (Calvert  Formation?)  than  from 
the  Cretaceous  Black  Creek  Formation. 

Eastman  (1899)  described  Amiopsis  dar- 
toni  from  a  partial  opercular  series,  pectoral 
fin,  and  associated  cycloid  scales  from  the 
Late  Jurassic  marine  Sundance  Formation, 
South  Dakota.  Eastman  felt  that  the  many 
"stout  ribs"  associated  with  the  pectoral  fin 
suggested  a  well-ossified  Aniia-hkc  verte- 
bral column  and  the  semicircular  operculum 
conformed  with  that  of  A.  ctilvu.  Since  the 
scalers  are  covered  supt^rficially  with  ganoine 
and  appear  elliptical,  Eastman  placed  this 
form  among  the  Amiidae.  He  allocated  the 
generic  name,  Amiopsis,  on  a  temporal 
basis.  According  to  Bobb  Schaeffer,  (1971, 
personal  communication)  the  holotype 
(USNM  4792)  and  the  paratypes  (MCZ 
9696,   USNM  4793)    are  to  be  tentatively 


referred  to  the  Leptolepidae  on  the  basis 
of  morphology  of  opercular  series  and  pec- 
toral fin  lepidotrichia.  Schaeffer  is  currently 
studying  the  Late  Jurassic  North  American 
fishes  and  is  including  a  more  extensive 
discussion  of  this  material  in  his  review. 

SUMMARY  AND  CONCLUSIONS 

This  survey  of  the  osteology,  morpho- 
metries, and  meristics  of  the  North  Amer- 
ican fossil  amiids  indicates  that  the  extant 
and  fossil  forms  fall  into  foiu-  groups 
worthy  of  specific  status:  ( 1)  Amia  fragosa, 
(2)  A.  uintaensis,  (3)  A.  scutata,  and  (4) 
A.  colvci.  All  these  forms,  excepting  A. 
fragosa,  have  somewhat  elongated  bodies 
(approximately  85  centra)  and  shai-p, 
conical  coronoid  and  palatal  teeth.  Al- 
though the  coronoid  and  palatal  teeth  of 
A.  uintaensis  are  more  sharply  curved  in- 
wardly, the  teeth  are  closer  in  morphology 
to  those  of  A.  scutata  and  A.  calva  than 
to  the  styliform  teeth  of  A.  fragosa.  A. 
uintaensis,  A.  fragosa,  and  A.  scutata  all 
have  a  larger  infraorbital  4  than  infraorbital 
5,  greater  degree  of  ossification  of  cranial 
elements,  deeper  orbital  notch  in  the  frontal, 
greater  head/ standard-length,  and  generally 
larger  parietal /frontal  ratio.  These  charac- 
ter-states clearly  set  the  fossil  species  of 
Amia  apart  from  the  Recent  A.  calva. 

Articulated  specimens  have  yielded  more 
information  on  the  osteology  of  A.  fragosa. 
A.  fragosa  is  a  short-bodied  form  (approxi- 
mately 65  centra)  with  a  smaller  number  of 
caudal  lepidotiichia  than  in  the  other 
species  of  Amia,  styliform  palatal  and  coro- 
noid teeth,  deeper  orbital  excavation  in  the 
frontals,  square  parietals,  and  a  short  box- 
like skull  having  relatively  short  mandibles 
diat  occupy  about  half  the  head-length.  The 
styliform  crushing  palatal  teetlr  of  A.  fragosa 
suggest  a  durophagous  habit,  rather  than 
the  more  predaceous  habit  indicated  by  the 
sharp  palatal  teeth  of  A.  uintaensis,  A. 
scutata,  and  A.  calva.  Although  it  is  known 
that  A.  calva  includes  molluscs  and  crusta- 
ceans in  its  diet,  perhaps  A.  fragosa  was 
more  exclusively  adapted  for  shell  crushing 
than  the  Recent  species. 


76         Bulletiti  Museuin  of  Comparative  Zoologtj,  Vol.   146,  No.   1 


Fig.  31.      Skull  and  body  structure  of  A,  Amia  calva;  B,  A.  scufafa;  C,  A.  uinfaensis;  and   D,  A.  fragosa. 


Fossil  Amiids  •  Boreske        77 


PLEISTOCENE 


PLIOCENE 


POST-BLANCAN 


BLANCAN- 


HEMPHILLIAN 


CLARENOONIAN 


FOSSIL  LAKE  BEOS 
(lOAHO  FM.) 


WAKEENEY   It. 
(OGALLALA  FM.) 

LOWER  VALENTINE    FM. 


BARSTOVIAN 


MIOCENE 


HEMINGFORDIAN 
ARIKAREEAN 


EUBANKS    l.f. 
(PAWNEE  CREEK   FM.) 


TURTLE  BUTTE  FM. 


WHITNEYAN 


OLIGOCENE 


ORELLAN 


CHAORONIAN 


RUBY  PAPER  SHALE 
(PASSAMARl    FM.) 


GRANT  HORSE  PRAIRIE  SHALE 

FLORISSANT    FM. 

ORELLA  MEMBER 
(BRULE   FM.) 


CHADRON   FM. 
CYPRESS  HILLS   FM. 


OUCHESNEAN 


EOCENE 


UINTAN 


BRIDGERIAN 


WASATCHIAN 


CLARNO  FM. 
HORSEFLY  RIVER  BEDS 


UINTA  FM. 
WASHAKIE  FM. 


BRIDGER  FM. 
WIND  RIVER  FM. 

FOSSIL  LAKE  BEOS 
(GREEN  RIVER  FM.) 

6OLOEN  VALLEY  FM. 

WASATCH  FM. 

GRAYBULL  BEDS 
(WILLWOOO  FM.) 


CLARKFORKIAN 


TIFFANIAN 


PALEOCENE 


TORREJONIAN 


PUERCAN 


MAASTRICHTIAN 


CRETACEOUS 


CAMPANIAN 


ALBIAN 


BEAR  CREEK  l.f, 
(FORT  UNION  FM.) 

SILVER  COULEE   l.f. 
(FORT  UNION  FM.) 


MELVILLE  FM. 
SAUNDERS  CREEK  l.f. 
(PASKAPOO  FM.) 

CEDAR  POINT  QUARRY  l.f. 
(FORT   UNION  FM  ) 


MEDICINE  ROCKS  l.f. 
(TONGUE  RIVER  FM.) 

ROCK  BENCH  l.f. 
(FORT  UNION  FM4 


TULLOCK  FM. 

MANTUA  If. 
(FORT  UNION  FM.) 


HELL  CREEK   FM. 
LANCE    FM. 
OJO  ALAMO  FM. 
AGUJA  FM. 
EDMONTON  FM. 


JUDITH  RIVER  FM. 
"MESAVERDE"  FM. 

OLDMAN  FM. 


BUTLER  FARM  l.f. 
(PALUXY  FM) 


7X 


Table  18.     Major  deposits  containing  remains  of  Amia  in  the 

WESTERN    interior    OF    THE    UnITEU    StATES    AND    CaNAUA 


78        Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Seven  genera  and  twenty-three  amiid 
species  (Table  19)  have  been  described  in 
the  literature.  Estes  (1964)  synonymized 
Stylomyleoclon  lacus  with  Kindleia  fra^osa, 
and  Estes  and  Berberian  ( 1969 )  referred 
the  genus  Kindleia  to  Amia,  thereby  con- 
firming the  suggestion  of  Janot  (1969). 
Paramiatus  p,iirleyi  (Romer  and  Fryxell, 
1928)  is  unquestionably  a  synonym  of  A. 
fragosa.  Regardless  of  possible  synonymy 
with  European  taxa,  the  sti'atigraphic 
range  of  A.  fragosa  is  remarkably  long, 
extending  as  it  does  from  the  Late  Creta- 
ceous through  the  Middle  Eocene.  Al- 
though A.  fragosa  is  better  known  than  the 
other  fossil  species,  and  was  extensively 
described  by  Estes  (1964),  O'Brien  (1969), 
and  Estes  and  Berberian  (1969),  its  phylo- 
genetic  relationship  to  them  and  to  A.  calva 
could  not  be  understood  without  compara- 
tive information  on  both  the  other  fossil 
forms  and  A.  calva  (Fig.  32). 

A.  newherrianus  and  A.  depressiis 
(Marsh,  1871),  and  A.  gracilis  (Leidy, 
1873a),  described  from  undiagnostic  ver- 
tebral characters,  are  considered  here  as 
nomina  duhia. 

A.  iiintaensis  is  a  form  having  a  relatively 
greater  body-length  than  the  other  species 
of  Amia.  It  has  approximately  the  same 
total  number  of  vertebrae  as  A.  calva  and 
A.  scutata,  but  the  arrangement  of  the 
coliunn  varies  meristically  from  them.  Its 
head  is  more  elongated  tlian  that  of  the 
other  forms,  with  the  jaws  occupying  over 
two-thirds  of  the  head-length.  The  vouier- 
ine  teeth  are  sharp  (as  are  the  palatal  and 
coronoid  teeth),  as  they  are  in  A.  scutata 
and  A.  calva,  but  are  more  than  twice  as 
numerous  as  in  these  later  forms.  The  pres- 
ent study  confirms  the  opinions  of  Romer 
and  Fryxell  ( 1928),  Estes  ( 1964),  and  Estes 
and  Berberian  (1969)  that  the  differences 
between  Amia  and  Protamia,  Hypamia,  and 
Pappichthys  are  insufficient  for  the  recogni- 
tion of  any  of  the  latter  as  genera  distinct 
from  Amia.  Hypamia  elegans  (Leidy, 
1873a)  is  considered  a  m)men  duhium,  be- 
ing based  on  vertebral  characters  that  can- 
not be  distinguished  from  those  of  the 
other    species.      Protamia    media     (Leidy, 


1873a),  Pappichthys  symphysis,  P.  corsonii, 
P.  medius,  P.  plicatus,  P.  sclerops,  P.  laevis 
(all  described  by  Cope,  1873),  as  well  as 
Atnia  macrospondyla  and  A.  whiteavesiana 
(Cope,  1891),  are  all  considered  here  as 
synonyms  of  A.  uintaensis;  they  were  based 
on  undiagnostic  vertebral  characters  and 
morphology  of  the  skull  elements.  Material 
of  large  amiids  from  the  Late  Cretaceous 
Lance  and  Hell  Creek  formations  is  referred 
to  A.  cf.  uintaensis,  since  the  material  differs 
only  in  minor  respects  from  the  Paleocene 
and  Eocene  specimens.  It  cannot  be  deter- 
mined whether  this  material  represents  ac- 
tual populations  of  A.  uintaensis  or  an 
earlier  stage  of  its  evolution.  The  strati- 
graphic  range  of  A.  uintaensis  extends  from 
the  Paleocene  to  the  Early  Oligocene. 

A.  scutata,  an  Early  to  Middle  Oligocene 
long-bodied  form,  shares  cranial  characters 
with  both  A.  uintaensis  and  A.  calva.  Al- 
though it  has  closer  morphometric  and 
meristic  affinities  to  the  Recent  form,  it  is 
structually  and  temporally  intermediate  be- 
tween A.  uintaensis  and  A.  calva;  it  resem- 
bles the  more  primitive  A.  uintaensis  in  the 
moi-phology  of  Meckel's  groove  and  coro- 
noid articulation  surface  of  the  dentary, 
greater  ossification,  and  in  having  an  elon- 
gated skull  with  a  greater  head/ standard- 
length  than  in  A.  calva.  A.  dictyocephala 
(Cope,  1875)  is  considered  a  synonym  of 
A.  scutata;  it  was  based  on  undiagnostic 
meristic  characters.  In  the  evolutionarv  con- 
tinuum,  A.  scutata  appears  to  be  an  inter- 
mediate stage  between  A.  uintaensis  and 
A.  calva  (Fig.  32).  A  more  direct  line  of 
evolution  exists  between  A.  scutata  and  A. 
calva;  this  is  supported  by  Miocene  and 
Pliocene  amiid  material  that  displays  cra- 
nial elements  closely  transitional  between 
the  two  species.  Thus  the  Recent  species  of 
A.  calva  had  begun  at  least  by  the  begin- 
ning of  the  Pliocene,  and  A.  calva  was  ap- 
parently distinct  from  A.  scutata  by  that 
time.  It  appears  that  A.  fragosa  represents 
an  amiid  population  that  survived  until  the 
Middle  or  Late  Eocene  and  had  no  phylo- 
genetic  affinities  with  the  modern  form  be- 
yond this  time. 


I 


Fossil  Amiids  •  Boreske        79 


/Im/'o  colva 

RECENT 

PLEISTOCENE 

POST-BLANCAN 

Rl    ANT  AM 

HEMPHILLIAN 

PLIOCENE 

CLARENDONIAN 

Ami  a  cf.  calva 

BARSTOVIAN 

A  mi  a  cf .  scut  at  a 

MIOCENE 

HEMINGFORDIAN 
ARIKAREEAN 

WHITNEYAN 

OLIGOCENE 

ORELLAN 
CHADRONIAN 

Ami  a  scuta  ta 
-       i 

DUCHESNEAN 

,     J 

EOCENE 

UINTAN 
BRIDGERIAN 

\ 

y 

WASATCHIAN 

^ragosa       Amia  uintaensis 

CLARKFORKIAN 

'V" 

/ 

PALEOCENE 

TIFFANIAN 

TORREJONIAN 

PUERCAN 

\ 

\  /     - 

MAASTRICHTIAN 

\      Amia  ct  uintaensis 

CRETACEOUS 

CAMPANIAN 

\  ; 

ALBIAN               Al 

niidae 

mcertae  sedi 

\ 

s 

Fig.   32.      Suggested  phylogenetic  relationships  within  the  genus  Am\a. 


80 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


In  the  North  American  fossil  record,  fossil 
remains  unquestionably  those  of  the  family 
Amiidae  first  occur  in  tlic  Lower  Cretaceous 
(Albian)  sediments  of  Texas.  However, 
none  of  the  material  can  be  referred  to  any 
known  species  of  Araia;  it  displays  charac- 
ter-states resembling  those  of  Amia  uintaen- 
sis  and  Amia  frcifi^osa,  as  well  as  the  Euro- 
pean Urocles.  Some  of  the  vertebrae  re- 
semble those  of  Amiopsis.  The  Paluxy  mate- 
rial may  represent  either  one  or  more  forms 
transitional  between  Amia  and  the  Late 
Mesozoic  European  Urocles  (or  Amiopsis) , 
or  an  as  yet  undescribed  line.  The  body- 
length  of  Amia  fra^osa  appears  to  be  a 
primitive  feature  derived  from  the  earlier 
amiids  Urocles,  Siruimia,  Ikechaoam,ia,  and 
Amiopsis.  Despite  thc>ir  different  vertebral 
columns,  Amia  frafi^osa  and  A.  uintaensis 
show  similar  morphology  of  the  cranial 
elements,  but  the  nature  of  the  probable 
common  origin  of  these  forms  is  still  uncer- 
tain in  the  absence  of  a  more  complete  fossil 
record. 

Remains  of  amiids  referable  to  or  close  to 
Amia  fragosa  and  A.  uintaensis  have  been 
described  from  the  Paleocene,  Eocene,  and 
Oligocene  of  Europe,  and  the  Eocene  of 
Asia.  Additional  but  still  not  definitive  evi- 
dence supports  Estes'  ( 1964 )  and  Estes  and 
Berberian's  (1969)  suggested  synonymy  of 
A.  russelU  (Late  Paleocene,  France),  A. 
kehreri  (Middle  Eocene,  Germany),  and 
A.  munieri  (Early  Oligocene,  France)  with 
A.  fragosa.  Pseudamia  Jieintzi  (Eocene, 
Spitzbergen )  and  A.  valenciennesi  ( Eocene, 
France)  are  also  possible  synonyms  of  A. 
fragosa.  A.  valenciennesi  is  the  oldest  name 
and  would  take  precedence  over  A.  fragosa. 
Cranial  similarities  confirm  the  synonymy  of 
A.  rohusta  (Late  Paleocene,  France)  with 
A.  uintaensis. 

European  and  North  American  fossil 
Amia  occurred  in  freshwater  deposits  and 
apparently  occupied  a  habitat  much  like 
that  of  the  Recent  species.  According  to 
Westoll  (1965:  19-20)  the  distribution  of 
freshwater  vertebrates  is  a  useful  indica- 
tion of  "direct  continental  communication," 


Table  19.     Amiid  genera  and  species  of  various 

AUTHORS  discussed  IN  TEXT  IN  RELATION  TO  THE 
REVISED   lAXONOMY 


Atnia  calva 

Kindlcia  fragosa 
Stijlomijleodon  lactis 
Amia  fragosa 
Paramiattts  f^tirlcyi 

Amia  scutata 
A7nia  dictyocepliala 
Aitiia  cxilis 

Protamia  uintaensis 
Protamia  media 
PappicJi th ys  m edius 
PappicJithys  pJicatus 
Pappichthys  sclerops 
Pappichthys  lacvis 
PappicJithys  symphysis 
PappicJithys  corsonii 
Atnia  loJiiteavcsiana 
Amia  macrospondyla 

Amia  depressus 
Amia  newJwrriamis 
Amia  gracilis 
Hypamia  elcf^ans 

Arnia  sehvyniana 


.  .  Amia  calva 
Amia  fragosa 

.Amia  sctitata 


Amia  uintaensis 


nomina  dubia 


.nomen  nudum 


since  ".  .  .  descendents  of  a  common  stock  on 
different  modern  continents  must  have  used 
essentially  a  terrestial  route."  The  present 
study  further  amplifies  similarities  in  the 
Paleocene  and  Early  Eocene  amiid  fossil 
record  of  North  America  and  Europe.  This 
distribution  of  amiids  adds  to  the  similarity 
of  assemblages  of  Paleocene  and  Early 
Eocene  lower  vertebrates  (Estes  et  al., 
1967)  and  mammals  (McKenna,  1972)  on 
the  two  continents.  The  occurrence  of 
Pseudamia  Iwintzi  in  the  Eocene  deposits 
of  Spitzbergen  may  be  additional  evidence 
for  the  existence  of  the  De  Geer  migration 
route  (bridging  Europe,  Spitzbergen,  and 
North  America  during  the  Paleocene  and 
until  the  close  of  Sparnacian  time),  espe- 
cially if  suggested  relationship  to  A.  fragosa 
could  be  demonstrated.  The  Asian  form  A. 
mongoliensis  resembles  A.  uintaensis  in 
minor  respects  but  is  sufficiently  distinct  in 
itself  to  be  maintained  as  a  separate  species. 


Fossil  Amiids  •  Boreske        8i 


LITERATURE  CITED 

Agassiz,  L.  1843.  Recherches  sur  les  poissons 
fossiles.    Neuchatel,  tomes  1-5,  atlases  1-5. 

Allis,  E.  1889.  The  anatomy  and  development 
of  the  lateral  line  system  of  Amia  calva.  J. 
Morphol.,  2:  1-540. 

.      1897.     The  cranial  muscles  and  cranial 

and  first  spinal  nerves  in  Amia  calva.  J.  Mor- 
phol., 12(3):   1-814. 

Ami,  H.  1891.  On  some  extinct  Vertebrata  from 
the  Miocene  rocks  of  the  north-west  Territories 
of  Canada  recently  described  by  Professor 
Cope.    Science,  18:  53. 

Andreae,  a.  1892.  Vorliiufige  Mitteilung  iiber 
die  Ganoiden  ( Lepidostetis  und  Amia)  des 
Mainzer  Beckens.  Verb.  Nat.  med.  Ver.  Hei- 
delberg (N.F.),  5:  7-15. 

.      1895.     Beitriige   zur   Kenntniss   der   fos- 

silen  Fische  des  Mainzer  Beckens.  Abh. 
Senck.  naturf.   Ges.,   18:  351-365. 

Becker,  H.  1961.  Oligocene  plants  from  the 
Upper  Ruby  River  Basin,  southwestern  Mon- 
tana. Mem.  Geol.  Soc.  Amer.,  82:  1-127. 

.     1962.     Two    new    species    of    Mahonia 

from  the  Grant-Horse  Prairie  Basin  in  south- 
western Montana.  Bull.  Torrey  Bot.  Club, 
89:  114-117. 

Blair,  W.,  A.  Blair,  P.  Brodkorb,  F.  Cagle, 
AND  G.  Moore.  1968.  Vertebrates  of  the 
United  States.  New  York:  McGraw-Hill 
Press,  450  pp. 

BoLK,  L.,  E.  Goppert,  E.  Kallius,  and  W.  Lu- 
BOSCH.  1936.  Handbuch  der  vergleichen- 
den  Anatomie  der  Wirbeltiere  4.  Berlin:  Ur- 
ban und  Schwarzenberg,   1116  pp. 

Boreske,  J.  1972.  Ta.xonomy  and  taphonomy  of 
the  North  American  fossil  amiid  fishes. 
(Abstr.)  Bull.  Geol.  Soc.  Amer.,  4(1):  3-4. 

Bridge,  T.  1877.  The  cranial  osteology  of  Amia 
calva.    J.  Anat.  Phys.,  11:  605-622. 

Cartier,  D.,  and  E.  Magnin.  1967.  La  crois- 
sance  en  longueur  et  en  poids  des  Atnia  calva 
L.  de  la  region  de  Montreal.  Canad.  J.  Zool., 
45:  797-804. 

Cavender,  T.  1968.  Freshwater  fish  remains 
from  the  Clarno  Formation  Ochoco  Moun- 
tains of  north-central  Oregon.  Ore  Bin,  30 
(7):   125-141. 

.     1970.     A    new   find   of   Amia   dictyoce- 

phala  Cope  from  the  Middle  Tertiary  Floris- 
sant Lake  Beds  of  western  North  America. 
Abst.  paper  given  at  50th  Annual  Meeting, 
Amer.  Soc.  Ichth.  Herp.:  42. 

Cope,  E.  1873.  On  the  extinct  Vertebrata  of 
the  Eocene  of  Wyoming,  observed  by  the  ex- 
pedition of  1872,  with  notes  on  the  geology. 
Ann.  Rept.  U.S.  Geol.  Surv.  Terr.  (1st  series), 
No.  6:  545-649. 

.      1875.     On    the    fishes    of    the    Tertiary 


shales  of  South  Park.  Bull.  U.S.  Geol.  Surv. 
Terr.  (2nd  series).  No.  1:  3-5. 

.      1884.     The   Vertebrata   of   the   Tertiary 

Formations  of  the  West.  Rept.  U.S.  Geol. 
Surv.  Terr.  (Hayden),  3:  745-746. 

.     1891.     On  Vertebrata  from  the  Tertiary 

and  Cretaceous  rocks  of  the  northwest  Terri- 
tory. Book  1.  The  species  from  the  Oligocene 
or  Lower  Miocene  of  Cypress  Hills.  Contrib. 
Geol.  Surv.  Canad.  Paleo.,  1:  2-4. 

Dean,  B.  1898.  On  the  dogfi.sh  (Amia  calva), 
its  habits  and  breeding.  Rept.  Comm.  Fish- 
eries, Game,  and  Forests  New  York,  No.  4: 
1-10. 

Dechaseaux,  C.  1937.  Le  genre  Amia,  son  his- 
toire  paleontologique.  Ann.  Paleont.,  26:  .3- 
16. 

Eastman,  C.  1899.  Jurassic  fishes  from  the 
Black  Hills  of  South  Dakota.  Bull.  Geol.  Soc. 
Amer.,  10:  397-407. 

Estes,  R.  1964.  Fossil  vertebrates  from  the  Late 
Cretaceous  Lance  Formation,  eastern  Wyo- 
ming. Univ.  Calif.  Publ.  Geol.  Sci.,  49:  1- 
180. 

,    AND    P.    Berberian.     1969.     Amia    (  = 

Kimlleia)  fragosa  (Jordan),  a  Cretaceous 
amiid  fish,  with  notes  on  related  European 
forms.  Breviora,  Mus.  Comp.  Zool.,  No.  329: 
1-14. 

, ,  AND  C.  Meszoely.     1969.     Lower 

vertebrates  from  the  Late  Cretaceous  Hell 
Creek  Formation,  McCone  County,  Montana. 
Breviora,  Mus.  Comp.  Zool.,  No.  337:  1-33. 

,    M.    HeCHT,    AND    R.    HOFFSTETTER.        1967. 

Paleocene   amphibians   from   Cernay,    France. 
Amer.  Mus.  Novitates,  No.  2295:   1-25. 
-,  AND  J.  TiHEN.      1964.     Lower  vertebrates 


from  the  Valentine  Formation  of  Nebraska. 
Amer.  Midi.  Nat.,  72(2):  453-472. 

Goodrich,  E.  1930.  Studies  on  the  Structure 
and  Development  of  Vertebrates.  London: 
Macmillan  Co.  Ltd.,  xxx  +  837  pp. 

Gould,  S.  1966.  Allometry  and  size  in  ontog- 
eny and  phylogeny.  Biol.  Reviews,  41:  587- 
640. 

Ha.mmett,  F.,  AND  R.  Hammett.  1939.  Pro- 
portional length  growth  of  gar  ( Lepisosteus 
platyrhincits  DeKay).  Growth,  3(2):  197- 
209. 

Hasse,  C.  1882.  Das  natiirliche  System  der 
Elasmobranchier.  Jena:  Verlag  von  Gu.stav 
Fischer,  pp.  77-284. 

Hatcher,  J.  1900.  The  Carnegie  Mu.seum  pale- 
ontological  expeditions  of  1900.  Science,  12 
(.306):   718-720. 

.  1901.  Some  new  and  little  known  fos- 
sil vertebrates.  Ann.  Carnegie  Mus.  Pitts- 
burgh, No.  1:   128-129. 

Hay,   O.     1895.     On  the  structure   and  develop- 


82         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


ment  of  the  vertebral  column  of  Amia.  Publ. 
Field  Columb.  Mus.  Zool.,  1(1):  1-54. 

.      1911.     Tlie      Pleistocene      of      Indiana. 

Ann.  Kept.  Ceol.  Nat.  Res.  Indiana,  .36:  552. 

.      1917.     Vertebrata   mostly   from    stratum 

No.  3,  at  Vero,  Florida;  together  with  de- 
scriptions of  new  species.  Rept.  Geol.  Surv. 
Florida,  9:  43-68. 

.      1923.     The  Pleistocene  of  North  Amer- 


ica and  its  vertebrated  animals  from  the  states 
east  of  the  Mississippi  River  and  from  the 
Canadian  Provinces  east  of  longitude  95  de- 
grees.   Bull.  Carnegie  Inst.,  .322:  336-382. 

HoFFSTETTER,  R.,  AND  J.  Gasc.  1969.  Verte- 
brae and  ribs  of  modem  reptiles.  In  Biology 
of  the  Reptilia.  New  York:  Academic  Press, 
xvi   -|-  468  pp. 

HuBBS,  C,  AND  K.  Lagler  1967.  Fishes  of  the 
Great  Lakes  Region.  Ann  Arbor:  University 
of  Michigan  Press,  viii  -|-  289  pp. 

HussAKOF,  L.  1932.  The  fossil  fishes  collected 
by  the  Central  Asiatic  Expeditions.  Amer. 
Mus.  Novitates,  No.  553:  1-16. 

Imbrie,  J.  1956.  Biometrical  methods  in  the 
study  of  invertebrate  fossils.  Bull.  Amer. 
Mus.  Nat.  Hist.,  108:  217-252. 

Janot,  C.  1966.  Aiuia  riisselli  nov.  sp.,  nouvel 
Amiide  ( Poisson  holosteen )  du  Thanetien  de 
Berru,  pres  de  Reims.  C.  R.  Soc.  Geol. 
France,  1966(3):   142. 

.      1967.     A  propos  des  amiides  actuels  et 

fossiles.  Colloq.  Intern.  C.  N.  R.  S.,  163: 
139-153. 

Jordan,  D.  1906.  The  fishes  of  Samoa.  Bull. 
Bureau  Fisheries,  25:  2.37. 

.      1919.     The  genera  of  fishes.    Publ.  Le- 

land  Stanford  Junior  University,  1919:  259. 

.  1925.  Opinion  89.  In  Opinions  ren- 
dered by  the  International  Commission  on 
Zoological  Nomenclature.  Smithsonian  Misc. 
Coll.,  73(3):  27-29. 

.      1927.     Kindleia,  a  new  genus  of  cichlid 

fishes  from  the  Upper  Cretaceous  of  Alberta. 
Canad.  Field  Nat.,  41:  145-147. 

.      1928.     Note   on   Kindleia   and   Stylomy- 

leodon  ( fossil  fish  from  Cretaceous  deposits 
of  Alberta).    Canad.  Field  Nat.,  42:  47. 

,  and  B.  Evermann.     1896.     The  fishes  of 


North  and  Middle  America.    Bull.   U.S.   Nat. 

Mus.,  47:  112-113. 
Lagler,  K.,  J.  Bardach,  and  R.  Miller.     1962. 

Ichthyology.     New    York:     John    Wiley    and 

Sons,  Inc.,  vii   +  523  pp. 
Lambe,    L.      1904.     Progress    in    vertebrate    pale- 
ontology in  Canada.   Trans.  Roy.  Soc.  Canada, 

No.  2:  27-43. 
.      1908.     The  Vertebrata  of  the  Oligocene 

of  the  Cypress  Hills,  Saskatchewan.    Contrib. 

Paleo.  Canada,  3(4):   12-13. 
Lange,     S.      1968.     Zur    Morphologic    imd    Tax- 

onomie  der  Fischgattung  Urocles  aus  Jura  nnd 


Kreide  Europas.  Palaeontographica,  Abt.  A, 
131:  1-78. 

Lehman,  J.  1951.  Un  nouvel  Amiide  de 
I'Eocene  du  Spitzberg,  Pseudamia  lieintzi. 
Troms0  Mus.  Arshefter,  70:  1-11. 

Leidy,  J.  1873a.  Notice  of  remains  of  fishes  in 
the  Bridger  Tertiary  Formation  of  Wyoming. 
Proc.  Acad.  Nat.  Sci.  Philadelphia,  1873: 
97-99. 

.      1873b.     Contributions     to     the     extinct 

vertebrate  fauna  of  the  western  territories. 
Rept.  U.S.  Geol.  Surv.  Terr.  (Hayden),  1: 
185-189. 

Liu,  T-S.,  H-T  Liu,  and  T-T  Su.  1963.  The 
discovery  of  Sinamia  zdanskyi  from  the  Ordos 
Region  and  its  stratigraphical  significance. 
Vert.  Palasiatica,  7(1):   1-30. 

Lund,  R.  1967.  An  analysis  of  the  propulsive 
mechanisms  of  fishes,  with  reference  to  some 
fossil  actinopterygians.  Ann.  Carnegie  Mus., 
39(15):   195-218. 

Marsh,  O.  1871.  Communication  on  some  new 
reptiles  and  fishes  from  the  Cretaceous  and 
Tertiary.  Proc.  Acad.  Nat.  Sci.  Philadelphia, 
1871:  103-105. 

McKenna,  M.  1972.  Was  Europe  connected 
directly  to  North  America  prior  to  the  Mid- 
dle Eocene?  In  Evolutionary  Biology,  Vol.  6. 
New  York:  Appleton-Century-Crofts,  pp.  179— 
189. 

Merrill,  G.  1907.  Catalogue  of  the  figured 
specimens  of  fossils,  minerals,  and  ores  in  the 
Department  of  Geology,  LTnited  States  Na- 
tional Museum.  Bull.  U.S.  Nat.  Mus.,  53: 
6-15. 

Miller,  H.  1968.  Additions  to  the  Upper  Cre- 
taceous vertebrate  fauna  of  Phoebus  Land- 
ing, North  Carolina.  J.  Elisha  Mitchell  Sci. 
Soc,  84:  467^71. 

Newton,  E.  1899.  On  the  remains  of  Amia, 
from  the  Oligocene  strata  in  the  Isle  of 
Wight.  Quart.  J.  Geol.  Soc.  London,  4:  1- 
10. 

Nybelin,  O.  1963.  Zur  Morphologic  und  Ter- 
minologie  des  Schwanzskelettes  der  Actinop- 
teiygier.    Arkiv.  Zool,  15(35):  485-516. 

O'Brien,  D.  1969.  Osteology  of  Kindleia  frag- 
osa  Jordan  (Holostei:  Amiidae)  from  the 
Edmonton  Formation  ( Maestrichtian )  of 
Alberta.  M.A.  Thesis,  University  of  Alberta: 
vi  -1-  118  pp. 

Osborn,  H.,  W.  Scott,  and  F.  Speir.  1878. 
Paleontological  report  of  the  Princeton  scien- 
tific expedition  of  1877.  Contrib.  Mus.  Geol. 
Arch.  Princeton,  No.  1:   102-104. 

Piton,  L.  1940.  Paleontologie  du  gisement 
Eocene  de  Menat  ( Puy-de-Dome,  flore  et 
faune).  Mem.  Soc.  Sci.  Nancy,  56:  1-303 
pp. 

Reighard,  J.     1903.     The  natural  history  of  Amia 


Fossil  A  muds  •  Bote  she 


83 


ralva    Linnaeus.     Mark    Anni\cisaiy    Vohinie 

(art.    t):  57-109. 
RoMKH,  A.,  AND  F.   FiiYXKLi,.      1 92S.     Purdiuiatiis 

^urlcyi,   a   ni-w   (Iccp-hocliccl   aiiiiid   fish    from 

tlie  Eocene  of  Wyoniinj,'.     Aiiiei.    |.    Sei.,    16 

(90):    519-527. 
|{us.st:LL,   L.      1928a.     A  new  fossil   fish    from   the 

Paskapoo  Beds  of  Alberta.     Aiiier.    |.   Sei.,    15 

(86):    103-107. 
.      19281).      The    jfenera    Kiiullcut    and    Sty- 

loviylcodon.    Anier.  J.  Sei.,   15(87):   264. 
.       1929.      Tiie    validity    of    the    .ueniis    Sttj- 


IniiujUoiUm.    Auier.  J.  Sei..  17(  100):  369-;571. 
.      1967.      Paleontology    of   the    Swan    Hills 

area,      north-central      Alheila.     Roy.      Ontario 

Mus.  C.'ontril).,  No.  71:    1-31. 
Sc:hakffeh,  B.      1967.      Osteichthyan  vertebrae.   |. 

Linn.  Soe.    (Zool.),  No.  47:    185-195. 
Shufixut,     R.      1885.     The     osteology    of    Ainiu 

calva.    Washington:  Government  Printing  Of- 
fice, 93  pp. 
SiMi'.soN,     (;.      1937.     The     Fort     Ihiion     of     the 

Crazy     Mountain     Field,     Montana,     and     its 

inannnalian    faunas.     Bull.     U.S.     Nat.     Mus., 

169:  \   +   287  pp. 
,     A.     RoK,     AND     R.     Lewontin.       1960. 

Quantitative   Zoology.     New   York:    Harcourt, 

Brace,  and  Co.,  iv   +  440  pp. 
Skinneh,  M.,  S.  Skinneh,  and  R.  Gooius.     1968. 

C-enozoic   rocks    and    faunas    of   Turtle    Butte, 

south-central  South  Dakota.    Bull.  Anier.  Mus. 

Nat.  Hist.,  138:  381-436. 
Smi'iii,     C.      1962.     Some     Pliocene     fishes     from 

Kansas,    Oklahoma,    and    Nebraska.     Copeia, 

1962(3):  505-520. 
Stensio,    E.     1935.     Smainia    zdanskyi,    a    new 


amiid  from  the  Lower  C^retaceoiis  of  Shan- 
tung, China.  Pahoiit.  Sinica,  .ser.  C,  No.  3: 
1-48. 

Swift,  C.  1968.  Fossil  bony  fishes  from  Flor- 
ida.   I'laster  Jacket,  No.  7:  2-11. 

SzALAY,  F.,  AND  M.  Mc:Kenna.  1971.  Begin- 
ning of  the  age  of  uiaiumals  in  Asia:  tin-  Late 
Paleocene  Cashato  fauna,  Mongolia.  Bvill. 
Amer.  Mus.  Nat.  Hist.,  144:  270-317. 

Thomson,  K.,  and  K.  IIaiin.  1968.  (Jrowth  ;nid 
form  in  fossil  rhipidistian  fishes  ( Cro.ssopter- 
ygii).    J.  Zool.,  156:   199-223. 

Thuhmond,  J.  1969.  Lower  vertebrates  and 
paleontology  of  the  Trinity  Croup.  Ph.D. 
Thesis,  Southern  Methodist  Unixersity  (  Libr. 
Congr.  No.  70-19259).  1.36  pp.  Univ.  micro. 
Ann  Arbor,  Mich,   (di.ss.  ab.str.  31:   2156). 

TnAQUAut,  R.  1911.  Les  poissons  wealdiens  de 
Bernissart.  Mem.  Mus.  Roy.  Hist.  Nat.  Belg., 
6:   1-65. 

Westoll,  T.  1965.  ecological  evidence  bear- 
ing upon  continental  drift.  Symposium  on 
Continental  Drift.  Royal  Soc.  London,  1965: 
12-25. 

WiUTEHousE,  R.  1910.  The  caudal  fin  of  the 
Teleostomi.  Proc.  Zool.  Soc.  London,  1910: 
590-627. 

Whitley,  C.  1954.  More  new  fish  names  and 
records.    Aust.  Zool.,  12:  57. 

Wilson,  R.  1968.  Systematics  and  faunal  anal- 
ysis of  a  Lower  Pliocene  vertebrate  assem- 
blage from  Trego  County,  Kansas.  Contril). 
Mus.  Paleo.   Univ.   Mich.,  22(7):   75-126. 

Woodward,  A.  1916.  The  fossil  fi.shes  of  the 
English  Wealden  and  Purbeck  formations. 
Palaeontogr.  Soc.  London,  Pt.  1,  1915:  1-48. 


84         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


5  cm 


afab. 


B 


.J<Vj, 


Plate    1.      A,    "Paramiatus   gurley'i"    FMNH    2201,    Early    Eocene,    Green    River    Formation,    Wyoming;    B,    Amia    iragosa 
MCZ  5347,  Early  Eocene,  Green  River  Formation,  Wyoming. 


Fossil  Amiids  •  Boreske        85 


Plate   2.     Amia   kebreri   BMNH    P33480,   collected    by   Walter   Kijhne   in    1951    from   Middle   Eocene   deposits   at  Messel 
bei  Darmstadt. 


86         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   1 


Plate  3.     Amia  uinfaensis  PU   13865,  Early  Eocene,  Green   River  Formation,  Wyoming. 


Fossil  Amiids  •  Bnrrslr 


?^"  ■_;.■■•«;>; 

m 

.:  '-^ 

-■ 

1    y 

•^fe%i 

'  • 

■•■-Tf^ 

^ 

•'i. 

-  .  i 

Plate   4.      Amio    scu/ofa.   Middle    Oligocene,    Florissant    Formation,    Colorado:    A,    counterpart    YPM    6243;    B,    counter- 
part USNM  4087;  C,  PU  10172. 


I 


i 


us  ISSN  0O27.4100 


Bulletin  OF  THE 

Museum   of 

Comparative 

Zoology 


An  Analysis  of  Variation  in  the  Hispaniolan 
Giant  Anole,  Anolis  ricordi  Dumeril 

and  Bibron 


ALBERT  SCHWARTZ 


HARVARD  UNIVERSITY 

CAMBRIDGE,  MASSACHUSETTS,  U.S.A. 


VOLUME  146,  NUMBER  2 
19  APRIL  1974 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED  BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 

Bulletin  1863- 

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Memoirs  1864-1938 

JoHNSONLV,  Department  of  MoUusks,  1941- 

OccAsioNAL  Papers  on  Mollusks,  1945- 

Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.    Fishes  of  the  Gulf  of  Maine. 
Reprint 

Brues,  C.  T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.    Classification  of 
Insects. 

Creighton,  W.  S.,  1950.    The  Ants  of  North  America.    Reprint. 

Lyman,  C.  P.,  and  A.  R.  Dawe  (eds.),  1960.    Symposium  on  Natural  Mam- 
malian Hibernation. 

Peters'  Check-list  of  Birds  of  the  World,  vols.  2-7,  9,  10,  12-15. 

Sprinkle,  J.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echinoderms. 

Turner,  R.  D.,  1966.    A  Survey  and  Illustrated  Catalogue  of  the  Teredinidae 
(MoUusca:  Bivalvia). 

Whittington,  H.  B.,  and  W.  D.  I.  Rolfe  (eds.),  1963.    Phylogeny  and  Evolu- 
tion of  Crustacea. 

Proceedings  of  the  New  England  Zoological  Club  1899-1948.     (Complete 
sets  only.) 

Publications  of  the  Boston  Society  of  Natural  History. 


Authors  preparing  manuscripts  for  the  Bulletin  of  the  Museum  of  Comparative 
Zoology  or  Breviora  should  send  for  the  current  Information  and  Instruction  Sheet, 
available  from  Editor,  Publications  OflBce,  Museum  of  Comparative  Zoology, 
Harvard  University,  Cambridge,  Massachusetts  02138,  U.S.A. 


©  The  President  and  Fellows  of  Harvard  College  1974 


AN  ANALYSIS  OF  VARIATION  IN  THE  HISPANIOLAN  GIANT 
ANOLE,  ANOLIS  RICORDI  DUMERIL  AND  BIBRON 


ALBERT  SCHWARTZ' 


Abstract.  The  nominal  Hispaniolan  species  of 
giant  anole,  Anolls  ricordi,  is  considered  to  be  in 
actuality  composed  of  three  distinct  allopatric  spe- 
cies: A.  ricordi,  A.  barahonae,  and  A.  baleatus. 
Subspecies  of  all  three  species  are  described,  but 
only  A.  baleatus  is  well  represented  in  collections. 
A  theoretical  history  of  this  species  complex  upon 
Hispaniola  is  presented. 

The  Hispaniolan  giant  anole,  Anolis  ri- 
cordi Dumeril  and  Bibron,  1837,  has  been 
known  to  science  for  more  than  a  century; 
yet  only  in  the  hist  35  years  has  it  become 
evident  that  this  species  is  not  homoge- 
neous in  its  characteristics  throughout  Haiti 
and  tlie  Republica  Dominicana.  The  spe- 
cies was  first  named  (as  Anolis  ricordii) 
from  Santo  Domingo,  as  the  entire  island 
was  known  at  that  historical  period,  but 
specimens  seem  to  have  been  rare  in  col- 
lections thereafter.  Schmidt  (1921:  10)  re- 
ported four  A.  ricordi  from  two  Dominican 
localities.  Cochran  (1941:  133)  Hsted  24 
specimens  (all  but  one  of  which  were  in 
the  National  Museum  of  Natural  History) 
from  11  localities.  Mertens  (1939:  68-70) 
studied  17  specimens  in  European  collec- 
tions and  was  the  first  to  recognize  that 
there  were  two  readily  distinguishable  pop- 
ulations that  he  considered  subspecies:  A. 
r.  ricordi  in  Haiti,  and  A.  r.  baleatus  Cope 
in  the  Republica  Dominicana.  Williams 
( 1962 )  reviewed  the  species  in  more  detail 
and  examined  90  specimens.  For  this  suite 
of   anoles,   he   described   A.    r.    barahonae 


\ 


1   Miami-Dade      Community      College,      Miami, 
Florida    33167. 

Bull.  Mus.  Co 


from  tlie  Sierra  de  Baoruco  in  the  south- 
western Republica  Dominicana.  Still  later, 
Williams  (1965)  studied  an  additional  80 
specimens  and  named  A.  r.  leberi  from 
Camp  Perrin  on  the  extreme  distal  portion 
of  the  Haitian  Tiburon  Peninsula.  Thus, 
with  increasing  quantities  of  material  from 
more  diverse  localities,  our  knowledge  of 
the  distribution  and  variation  in  this  species 
has  increased  accordingly. 

A  great  many  problems  remain,  however, 
when  one  deals  in  detail  with  the  variation 
in  A.  ricordi.  Williams  (1962,  1965) 
pointed  out  that  records  of  the  species  were 
of  such  a  scattered  nature  (especially  on 
the  Tiburon  Peninsula  but  also  elsewhere 
on  the  island)  that  intergrades  between 
several  of  the  subspecies  remained  unknown 
and  also  that  there  were  no  specimens 
available  from  large  areas  between  named 
populations.  Williams  and  Rand  (1969),  in 
their  excellent  summary  of  the  geographic 
differentiation  in  all  species  of  Hispaniolan 
anoles,  pointed  out  (p.  15)  that  Anolis  ri- 
cordi was  composed  of  "several  described 
subspecies,  some  of  which  are  sharply 
enough  distinct  to  raise  the  question  of  pos- 
sible species  status."  This  is  most  especially 
true  of  the  taxa  ricordi,  baleatus,  and  bara- 
honae, all  of  which  are  extremely  well 
characterized  by  both  pigmental  and  struc- 
tural details,  but  all  of  which  occupy  areas 
(extensive  in  the  cases  of  ricordi  and  ba- 
leatus) without  known  intergradation  be- 
tween them  or  without  close  geographic 
approximation.    Thus,      the      closest      ap- 

mp.  Zool.,   146(2):   89-146,  April,   1974  89 


90 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


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HiSPANiOLAN  Giant  Angle  •  Schwartz         91 


proaclu's  of  the  ranges  of  ricordi  and  1)0-  material  upon  whieh  A.  r.  leheri  was  based. 

leatits    (Copey    and    Peiia,    both    in    the  Our  unique  experience  at  Camp  Perrin — 

Republica   Dominieana)    are  separated  by  namely,  of  many  A.  ricordi  received  from 

some   115  kilometers  airline.    The  subspe-  local    Haitians — showed    that    the    species 

cies  harahomie  and  ricordi  (Sierra  de  Bao-  may  not  be  necessarily  rare.    However,  as 

rueo   and   associated   eastern   coastal   areas  in  A.  equestris  in  Cuba,  the  cryptic  greens 

in  the  Republica  Dominieana,  and  Saltrou  of  A.  ricordi  render  the  species  virtually  in- 

in  Haiti)   are  known  from  localities  sepa-  visible  during  the  day  except  to  all  but  the 

rated  by  about  115  kilometers,   and  bara-  most  experienced  observer.    In  1963,  Rich- 

honae  and  balcatus  by  a  gap  of  about  115  ard    Thomas    discovered    that    A.    ricordi 

kilometers  (between  the  Sierra  de  Baoruco  might  be  seemed  at  night,  since  individ- 

and  near  Villa  Altagracia,  both  in  the  Re-  uals  sleep  quite  exposed  in  a  variety  of  ar- 

publica  Dominieana).  boreal  situations  and  are  very  conspicuous. 

Schwartz  and  Garrido  (1972)  recently  Thus,  with  the  knowledge  that  the  agroma 
showed  that  the  Cuban  giant  anole,  Anolis  (as  the  species  is  known  in  Haitian  Creole) 
equestris  Merrem,  is,  in  fact,  a  complex  of  or  the  saltacocote  ( as  the  species  is  known 
five  species;  they  also  suggested  (p.  71)  in  Dominican  Spanish)  might  be  common 
that,  as  Williams  and  Rand  had  pointed  and  thus  easily  secured  by  native  collectors, 
out,  there  was  a  good  possibility  that  the  and  that  individuals  might  be  readily  se- 
Hispaniolan  Anolis  ricordi  in  time  might  cured  at  night  while  they  slept,  I  had  as 
likewise  be  shown  to  be  a  complex  of  spe-  one  of  my  objectives  to  secure  as  many  A. 
cies.  It  is  the  purpose  of  the  present  paper  ricordi  as  possible  in  order  to  clarify  the 
to  discuss  the  variation  in  A.  ricordi,  based  status  of  the  named  subspecies  and  in  an 
upon  the  examination  of  403  specimens  attempt  to  narrow  the  geographic  gaps  that 
from  a  broad  selection  of  geographic  local-  seemed  to  exist  between  ricordi,  baleatus, 
ities  throughout  Hispaniola.  Despite  my  and  barahonae.  As  more  material  accumu- 
ha\'ing  studied  far  more  material  than  any  lated,  we  were  successful  in  the  latter  at- 
previous  investigator,  there  still  remain  tempt,  but  the  range  of  variation  in  newly 
many  problems  that  cry  out  for  solution,  acquired  material  showed  that  the  situation 
As  Schwartz  and  Garrido  also  pointed  out  was  more  complex  than  was  supposed.  In 
in  their  analysis  of  Anolis  equestris,  the  addition  to  specimens  in  the  Albert 
present  paper  in  no  way  should  be  consid-  Schwartz  Field  Series  (ASFS),  collected 
ered  as  the  final  word  on  A.  ricordi;  rather  by  myself  and  field  assistants,  I  have  ex- 
it is  an  attempt  to  comment  in  detail  upon  amined  material  in  the  American  Museum 
the  known  variation  and  distribution  of  of  Natural  History  (AMNH),  the  Museum 
this  species  in  Hispaniola  which  may  serve  of  Comparative  Zoology  (MCZ),  and  the 
as  a  stepping  stone  for  further  work  upon  National  Museum  of  Natural  History 
the  species.  (  USNM).   For  the  loans  of  specimens  I  am 

Between  1962  and  1971,  I  and  my  asso-  grateful  to  Richard  G.  Zweifel,  George  W. 

ciates   collected   extensively  in  both   Haiti  Foley,  Ernest  E.  Williams,  and  George  R. 

and  the  Republica  Dominieana.    Latterly,  Zug.   In  all   of  these  collections   there  are 

between  1968  and  1971,  my  work  in  His-  other  specimens   that   I    have   deliberately 

paniola  has  been  under  the  sponsorship  of  not  elected  to  study,  since  many  of  them 

two   National    Science    Foundation    grants,  are  from  localities  that  are  now  well  repre- 

GB-7977  and  B-023603.    Specimens  of  Ano-  sented  by  more  recently  collected  lizards 

lis  ricordi  collected  in  1962-63  were  avail-  or  that  have  poor  locality  data.    Specimens 

able  to  Williams  and  were  reported  upon  in  the  collection  of  the  Museum  of  Com- 

by  him  (1965);  in  fact,  the  long  series  of  parative  Zoology  have  been  collected  un- 

A.  ricordi  from  Camp  Perrin,  Haiti,  secured  der    NSF    grant    B-019(S01X    and    previous 

for  me  by  native  collectors  in  1962,  was  the  grants   to   Dr.   Williams.     Most   of  the  re- 


92         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


cently  taken  ASFS  A.  ricordi  have  detailed 
descriptions  of  color  and  pattern  in  life, 
but,  as  in  all  such  endeavors  that  span  sev- 
eral years,  it  is  unfortimate  that  all  details 
of  color  and  pattern  have  not  been  consis- 
tently recorded  as  time  has  passed.  Like- 
wise, there  are  no  color  or  pattern  data  on 
most  old  specimens;  thus,  I  feel  less  secure 
in  dealing  with  these  older  specimens  or 
those  collected  by  others  than  myself  and 
parties  than  I  am  with  those  in  the  ASFS 
which  are  carefully  documented.  However, 
specimens  from  other  localities  must  in 
some  way  be  dealt  with,  and  I  have  done 
so  as  carefully  as  possible,  considering  de- 
tails of  geography  and  what  is  known  about 
specimens  of  A.  ricordi  from  adjacent  local- 
ities. 

I  wish  to  acknowledge  with  enthusiasm 
the  efforts  on  my  behalf  in  the  field  of  the 
following  men,  without  whose  efforts  the 
quantity  of  A.  ricordi  presently  available 
to  me  would  be  far  less:  Jeffrey  R.  Buffett, 
Carl  Butterfield,  James  R.  Dennis,  Danny 
C.  Fowler,  Ronald  F.  Klinikowski,  David 
C.  Leber,  James  A.  Rodgers,  Jr.,  Bruce  R. 
Sheplan,  and  Richard  Thomas.  C.  Rhea 
Warren  has  given  me  a  specimen  of  A.  ri- 
cordi from  northern  Haiti.  My  notes  on 
coloration  and  pattern  of  A.  ricordi  have 
been  greatly  supplemented  during  the 
present  study  by  the  color  portraits  exe- 
cuted in  the  field  by  D.  C.  Leber;  one  of 
these  has  been  reproduced  in  black-and- 
white  in  Williams  ( 1965 ) ,  but  the  repro- 
duction hardly  does  justice  to  the  detailed 
beauty  of  all  the  originals.  I  have  been  able 
to  examine  the  holotype  of  Eupristis  ba- 
leattis  Cope  through  the  courtesy  of  Alice 
C.  C.  Grandison  and  A.  F.  Stimpson  of  the 
British  Museum  (Natural  History).  Holo- 
types  and  paratypes  have  been  designated 
or  deposited  in  the  above  collections  and 
in  the  Caniegie  Museum  (CM). 

THE  PROBLEM 

Mertens  ( 1939 )  was  the  first  to  point  out 
that  Haitian  and  Dominican  A.  ricordi  dif- 
fered from  each  other  in  one  notable  char- 
acter— the  height  of  the  dorsal  crest  scales. 


His  figure  41  shows  this  character  ex- 
tremely clearly:  in  nominate  ricordi  from 
Haiti,  the  nuchal  crest  scales  are  low  and 
inconspicuous,  whereas  in  Dominican  ba- 
leatiis  the  nuchal  crest  scales  are  long  and 
attenuate.  In  addition,  Mertens  (1939:  69) 
characterized  ricordi  as  having  9  to  12 
scales  between  the  eyes;  males  of  this  sub- 
species have  one  or  more  sharply  defined 
black  blotches  on  the  sides  of  the  nape,  the 
occipital  area  flecked  with  black,  and  often 
have  black  longitudinal  stripes  on  the 
flanks.  On  the  other  hand,  baleatus  has 
from  6  to  8  scales  between  the  eyes,  and 
males  are  without  any  black  head,  nape,  or 
lateral  markings.  Williams  (1962)  com- 
pared these  two  taxa  with  barahonae  in  re- 
gard to  four  characters:  height  and  relative 
length  of  nuchal  and  dorsal  crest  scales, 
number  of  snout  scales  at  the  level  of  the 
second  canthal  scale,  and  body  pattern. 
Later,  when  he  described  A.  r.  leberi,  Wil- 
liams (1965)  employed  these  same  charac- 
ters to  differentiate  that  subspecies. 

The  differences  in  these  characters  be- 
tween the  four  recognized  subspecies  are 
unequivocal:  one  can  differentiate  at  a 
glance  between  such  distinctive  animals  as 
leberi  and  barahonae  or  between  ricordi 
and  baleattis,  without  recourse  to  micro- 
scopic examination.  The  whole  a.spects  of 
all  four  taxa  are  quite  distinctive,  whether 
one  is  dealing  with  living  or  long-preserved 
animals.  What  has  been  equivocal  is  the 
relationships  of  these  four  taxa,  since,  as  I 
pointed  out  previovisly,  they  have  been 
known  from  rather  isolated  groups  of  lo- 
calities, widely  separated  from  each  other. 
In  only  one  case  (leberi-ricordi)  have 
specimens  been  regarded  as  intergradient 
between  two  subspecies:  these  intergrades 
are  from  a  geographically  plausible  locality 
that  itself  is  widely  removed  from  the  two 
"parent"  populations. 

As  material  has  gradually  accumulated, 
it  has  become  increasingly  obvious  that  the 
situation  is  even  more  complex  than  has 
been  previously  recognized.  For  example, 
in  1971,  I  had  occasion  to  compare  long  se- 
ries of  living  examples  from  the  Peninsula 


HisPANioLAN  Giant  Anoll.  •   Schwuiu 


de  Sainana  and  the  adjacent  "mainland"  at 
Cafio  Abajo,  and  I  was  at  once  struck  with 
the  differences  between  tliese  two  samples, 
both  of  which  have  been  regarded  as  ha- 
leattis.  In  this  case,  the  differences  are  not 
particularly  subtle  but  they  do  involve  dif- 
ferences in  coloration  and  pattern  which 
often  are  evanescent  after  preservation. 
The  same  statement  may  be  made  about  A. 
ricordi  from  the  region  near  La  Vega  and 
those  from  the  Cordillera  Septentrional.  In 
1971  I  had  occasion  to  collect  specimens 
from  both  these  regions  on  two  successive 
days  and  thus  was  able  to  compare  freshly 
collected  material  directly.  Again,  the  dif- 
ferences are  ones  of  pattern  and  color,  but 
they  are  so  striking  that  it  is  misleading  to 
consider  both  these  populations  as  being 
identical  genetically.  I  could  multiply  the 
above  examples  but  to  no  purpose;  it  is  ob- 
vious, when  one  sees  living  A.  ricordi  in  the 
field,  that  there  are  several  populations 
presently  assigned  to  haleatus  which  are 
quite  distinctive. 

On  the  basis  of  specimens  collected  by 
Richard  Thomas  and  myself  in  1963,  Wil- 
liams ( 1965 )  reported  A.  r.  ricordi  for  the 
first  time  from  the  northwestern  Republica 
Dominicana  in  the  region  near  Pepillo  Sal- 
cedo  and  Copey  in  Monte  Cristi  Province. 
He  noted,  however,  that,  "Despite  the  new 
collections  one  embarrassment  remains.  No 
certain  intergrades  between  the  two  strik- 
ingly different  forms  ricordii  and  ])aleatiis 
are  yet  known.  .  .  .  However,  the  area  in 
which  intergrades  may  occur  is  being  nar- 
rowed: on  the  north  coast  of  the  Dominican 
Republic  between  Monte  Cristi  and  Santi- 
ago and  in  the  center  of  Hispaniola  be- 
tween Mirebalais  (  MCZ  68479,  69404)  and 
Santiago.  This  still  leaves  a  very  wide  area 
of  ignorance."  Since  the  above  was  written, 
I  have  secured  specimens  of  the  nominate 
subspecies  in  four  other  regions:  at  Re- 
stauracion,  Dajabon  Province,  along  the 
Dominico-Haitian  border  and  about  40  ki- 
lometers airline  south  of  the  Monte  Cristi 
localities;  on  the  southern  slopes  of  the  Cor- 
dillera Central  near  Juan  de  Herrera  in  San 
Juan  Province;  and  throughout  the  Sierra 


de  Neiba  between  Hondo  Valle  and  Valle- 
juelo  in  La  Lstrelleta  and  San  Juan  prov- 
inces. These  latter  two  regions  (the  south- 
ern slopes  of  the  Cordillera  Central  and  the 
Sierra  de  Neiba)  are  separated  by  the 
rather  xeric  Valle  de  San  Juan.  Elsewhere 
to  the  east  and  north,  the  Cordillera  Cen- 
tral harbors  A.  ricordi  with  long  nuchal 
crest  scales  and  without  black  nape  and 
head  markings  in  males  (i.e.,  —  haleatua), 
as  at  San  Jose  de  Ocoa,  La  Vega,  and  the 
interior  uplands  near  El  Rio,  and  near  Los 
Montones  on  the  Rio  Bao.  The  fourth  lo- 
cality is  perhaps  the  most  significant;  there 
is  one  subadult  male  from  Santiago  Ro- 
driguez Province  near  Los  Quemados  which 
is  clearly  a  ricordi.  Of  the  haleatus  locali- 
ties, this  one  is  closest  to  Los  Montones,  a 
distance  of  50  kilometers  airline.  Thus  the 
gap  between  ricordi  and  haleatus  in  north- 
ern Republica  Dominicana  has  been  more 
than  halved,  and  there  still  is  no  genetic  in- 
fluence of  one  subspecies  upon  the  other. 

To  the  south,  specimens  of  A.  ricordi 
from  the  Sierra  Martin  Garcia,  and  Azua 
and  Peravia  provinces  likewise  narrow  the 
gap  there  between  haleatus  and  ricordi  on 
one  hand  and  between  haleatus  and  hara- 
honae  on  the  other.  In  the  former  case,  the 
distance  between  ricordi  and  haleatus 
(Vallejuelo  and  Sierra  Martin  Garcia)  is 
about  60  kilometers  airline,  without  char- 
acter dilution.  In  the  instance  of  harahonae 
and  haleatus,  only  20  kilometers  separate 
known  localities  (Barahona  and  Sierra 
Martin  Garcia)  for  these  two  taxa:  how- 
ever, the  actual  kilometrage  is  deceiving, 
since,  lying  between  these  localities,  is  the 
Valle  de  Neiba  and  the  Bahia  de  Barahona. 
Although  this  eastern  extreme  of  the  Valle 
de  Neiba  is  rather  mesic  and  supports  (or 
did  support )  large  trees  in  many  areas  that 
would  presumably  be  suitable  for  A.  ri- 
cordi, the  break  between  these  two  popu- 
lations is  sharp  despite  presumably  suitable 
ecology.  I  have  little  doubt  that  A.  ricordi 
occurs  in  this  intervening  region,  and  the 
interaction  of  harahonae  and  haleatus 
therein  will  be  most  interesting  to  ascertain. 

Even  more  intriguing  is  the  fact  that  the 


94         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


Sierra  Martin  Garcia  is  an  extreme  eastern 
isolate  of  the  Sierra  de  Neiba,  which  is 
elsewhere  occupied  by  A.  r.  ricordi.  This 
small  range,  which  reaches  an  elevation  of 
1350  meters,  is  completely  surrounded  by 
extremely  arid  desert  or  low  rolling  xeric 
hills,  as  well  as  by  the  Bahia  de  Barahona 
on  its  southwestern  edges.  The  heipeto- 
fauna  of  the  Sierra  Martin  Garcia  is  just 
becoming  knowti,  and  it  supports  a  remark- 
able fauna,  including  an  endemic  species 
of  Diplog,Iossus  (Thomas,  1971)  and  a  new 
species  of  Sphaerodactylus,  as  well  as  other 
unexpected  novelties.  Nevertheless,  A.  ri- 
cordi seems  to  have  reached  this  range 
from  the  northeast  (i.e.,  the  southern  slopes 
of  the  Cordillera  Central),  since  I  regard 
the  Martin  Garcia  lizards  as  identical  to 
those  from  Azua  and  Peravia  provinces. 

Finally,  the  geographic  relationship  of 
barohonae  and  ricordi  has  been  to  some  ex- 
tent clarified.  A.  r.  harahonae  has  been 
known  only  from  the  eastern  uplands  of 
the  Sierra  de  Baoruco  and  from  three 
southern  lowland  localities  ( Enriquillo, 
half-way  between  Enriquillo  and  Oviedo, 
and  Oviedo).  Each  of  the  latter  localities 
is  represented  by  a  single  specimen.  The 
Enriquillo  and  Enriquillo-Oviedo  speci- 
mens are  quite  obviously  harahonae,  but, 
as  Williams  (1965:  4)  noted,  the  specimen 
from  close  to  Oviedo  is  quite  different  in 
style  of  pattern  and  color  from  typical 
harahonae.  To  the  west,  in  Haiti,  there  has 
been  but  a  single  specimen  from  Saltrou 
which  Williams  (1965:  2)  considered  A.  r. 
ricordi  and  which  "narrows  the  geographic 
gap  between  ricordii  ricordii  and  r.  hara- 
honae; however,  it  does  nothing  to  narrow 
the  character  gap."  Two  additional  facts 
are  important.  First,  in  1971,  we  secured 
a  pair  of  A.  ricordi  from  along  the  Domin- 
ico-Haitian  border  north  of  Pedernales; 
these  individuals,  although  differing  some- 
what from  typical  harahonae  and  strongly 
from  the  single  Oviedo  specimen,  in  no 
way  show  any  tendencies  toward  A.  r.  ri- 
cordi. They  are  clearly  related  to  hara- 
honae, a  rather  surprising  fact  since  they 
are  much  closer  (35  kilometers)  to  Saltrou 


than  they  are  to  any  harahonae  locality  (65 
kilometers  at  Enriquillo).  Secondly,  Wil- 
liams has  recently  received  a  fine  se- 
ries of  A.  ricordi  from  Source  Carroye 
near  Thiotte;  these  lizards  are  quite  obvi- 
ously not  A.  r.  ricordi  but  are  closer  in 
many  ways  to  the  far-western  A.  r.  leheri. 
Thus  the  situation  along  the  southern  Hai- 
tian coast  between  Saltiou  and  the  eastern 
coast  of  the  Republica  Dominicana  at  Ovi- 
edo and  its  environs  remains  a  true  puzzle. 
It  seems  likely  that  A.  r.  ricordi  does  not 
cross  the  high  Massif  de  la  Selle,  except 
possibly  by  some  circuitous  route,  and  that 
A.  r.  harahonae  occurs  up  to  the  Dominico- 
Haitian  border,  to  within  11  kilometers  of 
a  station  ( Source  Carroye )  where  another 
taxon  occurs,  without  character  dilution. 

Interpretations  of  all  these  facts  are  seri- 
ously hampered  by  the  lack  of  specimens  of 
A.  ricordi  from  throughout  the  Haitian  Ti- 
buron  Peninsula.  Material  from  the  penin- 
sula may  be  divided  into  four  basic  lots:  a 
short  series  from  the  region  about  Castillon 
on  the  northern  slopes  of  the  Massif  de  la 
Hotte  near  the  tip  of  the  peninsula;  a  very 
long  series  of  specimens  from  Camp  Perrin 
on  the  low  southern  slopes  of  the  Massif  de 
la  Hotte  (the  type  series  of  A.  r.  leheri); 
a  short  series  from  midway  along  the  pen- 
insula at  Miragoane-Paillant;  and  a  moder- 
ate number  of  specimens  from  near  the 
base  of  the  peninsula  in  the  region 
of  Morne  Decayette-Petionville-Port-au- 
Prince.  The  lack  of  material  from  such 
well-collected  areas  as  Jeremie  on  the 
northwestern  coast  or  Jacmel  and  Les 
Cayes  on  the  southern  coast  is  extremely 
puzzling — we  simply  know  nothing  about 
lowland  A.  ricordi  throughout  much  of  the 
Tiburon  Peninsula,  except  for  the  above 
scattered  records  and  the  southern  coast  at 
Saltrou  near  the  Dominican  border  (and 
the  latter  locality  is  not  even  on  the  penin- 
sula proper). 

To  summarize  the  data  from  elsewhere 
in  Haiti  and  the  Republica  Dominicana, 
there  is  a  huge  distributional  hole  in  cen- 
tral Haiti,  with  but  two  specimens  {ri- 
cordi)   from   Mirebalais,   widely  separated 


HisPANiOLAN  Giant  Anoli 


b.:,/Ui   i: 


from  southern  ricordi  at  and  near  Port-an- 
Prince,  and  then  a  group  of  seattered  local- 
ities along  the  northern  littoral  of  Haiti 
from  Port-de-Paix  in  the  west  to  Terrier 
Rouge  in  the  east,  and  one  specimen  from 
Marmelade  in  the  interior  Chaine  de  Mar- 
melade.  All  these  Haitian  specimens  have 
low  nuchal  crest  scales,  and  males  variably 
possess  some  black  nape  and  side  markings, 
but  there  are  differences  between  speci- 
mens from  the  various  sections  which  pres- 
ently defy  analysis,  since  the  material  is 
too  meagre  and  from  too  scattered  locali- 
ties. 

The  Republica  Dominicana  fares  far  bet- 
ter as  far  as  detailed  coverage  is  con- 
cerned. Aside  from  the  material  previously 
noted  as  assigned  to  ricordi  or  harahonae, 
there  are  now  good  series  available  from 
the  eastern  half  of  the  country,  and,  al- 
though there  are  certain  gaps  even  within 
this  region,  they  are  not  so  appalling  as  are 
those  in  Haiti.  A.  ricordi  is  rarer  (or  per- 
haps less  easily  encountered)  in  arid  re- 
gions, and  thus  the  most  striking  gaps  in 
the  western  half  of  the  Republica  Domini- 
cana are  those  involving  arid  regions  on 
the  one  hand  or  high  mountain  masses  on 
the  other.  The  latter  situation,  especially  if 
the  slopes  are  pine-clad,  does  not  appear 
suitable  for  A.  ricordi.  and  the  species  may 
be  truly  absent  from  the  uplands  above 
4000  feet  (1220  meters),  the  highest  eleva- 
tion from  which  the  species  is  known.  In 
arid  regions,  A.  ricordi  appears  to  be  re- 
stricted to  riverine  woods  and  forests;  in 
such  situations,  the  species  may  not  be  un- 
common, but  it  may  require  diligence  to 
secure  even  one  specimen  from  a  particular 
region. 

The  detailed  discussion  above  should 
give  the  reader  a  background  of  both  the 
history  and  present  knowledge  of  the  dis- 
tribution of  A.  ricordi  against  which  the 
following  accounts  can  be  most  logically 
followed.  One  further  point  is  of  interest. 
A.  ricordi  is  unknown  by  specimens  from 
any  of  the  large  Hispaniolan  satellite  is- 
lands. In  some  cases  (Isla  Beata)  the  spe- 
cies is  not  expected  for  a  variety  of  reasons, 


but  in  others  (He  de  la  Gonave,  lie  dc  la 
Tortuc,  Ile-a-Vache)  there  seems  no  logical 
reason  for  the  absence  of  A.  ricordi,  dis- 
counting the  vagaries  of  overseas  transpoit. 
The  species  does  occur  on  Isla  Saona,  but 
remains  uncollected  there.  Fowler  and 
Sheplan  saw  a  sleeping  A.  ricordi  on  the 
northern  coast  of  Isla  Saona  in  December 
1971,  but,  after  it  had  been  shot,  it  was 
lost  in  the  undergrowth.  The  occurrence  of 
A.  ricordi  on  any  Hispaniolan  satellite  is 
noteworthy,  and  it  will  be  most  interesting 
to  determine  the  status  of  the  Isla  Saona 
population. 

METHODOLOGY 

The  series  of  403  A.  ricordi  was  divided 
into  14  samples  on  the  basis  of  geography, 
as  follows:  Republica  Dominicana:  1)  Pe- 
ninsula de  Samana  (54  specimens);  2) 
northeastern  Republica  Dominicana,  from 
Duarte  and  eastern  La  Vega  provinces  east 
to  the  haitises  region  in  northeastern  San 
Cristobal  Province  ( 37 ) ;  3 )  extreme  east- 
ern Hispaniola,  Punta  Cana-Juanillo,  Boca 
de  Yuma,  La  Altagracia  Province  ( 16 ) ;  4 ) 
southeastern  Republica  Dominicana  from 
Higiiey  and  Las  Lisas,  La  Altagracia  Prov- 
ince, west  to  Santo  Domingo  and  Yamasa, 
San  Cristobal  Province  (61);  5)  lowlands  at 
the  northern  base  of  the  Cordillera  Central 
at  Guaigui,  La  Vega  Province  (21);  6) 
Cordillera  Central  at  and  above  elevations 
of  2000  feet  (18);  7)  Cordillera  Septentri- 
onal and  north  ( 15 ) ;  8 )  Sierra  Martin  Gar- 
cia and  Peravia  and  Azua  provinces  (6);  9) 
Sierra  de  Baoruco  and  associated  east  coast 
of  the  Peninsula  de  Barahona  (33);  10) 
Oviedo,  Pedemales  Province  ( 1 ) ;  Haiti: 
11)  Saltrou  and  vicinity,  Dept.  de  I'Ouest 
( 15 ) ;  12 )  region  about  Port-au-Prince, 
Mirebalais,  northern  Haitian  littoral, 
Chaine  de  Marmelade,  and  (in  the  Repu- 
blica Dominicana)  region  about  Pepillo  Sal- 
cedo,  Copey,  Los  Quemados,  Restauracion, 
and  the  southern  slopes  of  the  Cordillera 
Central  and  the  Sierra  de  Neiba  (50);  13) 
Camp  Perrin  and  Marceline,  Dept.  du  Sud 
(54);  and  14)  vicinity  of  Castillon,  Dept. 
du  Sud  (6).   This  division  into  14  regional 


96 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


samples  was  completed  after  preliminary 
examination  of  the  material,  scale  counts 
and  detailed  review  of  field  notes  on  color- 
ation and  pattern  were  all  made.  In  addi- 
tion, two  other  small  lots  (eight  specimens 
from  the  vicinity  of  Miragoane,  Dept.  du 
Sud,  Haiti,  and  seven  specimens  from  El 
Seibo  Province,  Republica  Dominicana) 
were  examined  separately;  in  both  cases, 
these  short  series  indicate  intergradient 
tendencies  between  pairs  of  the  14  major 
samples  noted  above,  and  they  were  not  in- 
cluded with  the  latter. 

The  following  data  were  taken  on  each 
specimen: 

1 )  Snout-vent  length,  in  millimeters. 

2)  Number  of  snout  scales  across  snout 
at  level  of  the  second  canthal  scale,  reck- 
oned from  the  anterior  corner  of  the  orbit. 

3)  Number  of  vertical  rows  of  loreal 
scales. 

4)  Minimum  number  of  scales  between 
supraorbital  semicircles. 

5)  Number  of  scales  between  the  inter- 
parietal scale  and  the  supraorbital  semicir- 
cles on  each  side,  this  datum  written  as  a 
fraction  ( i.e.,  5/5  =  five  scales  in  this  posi- 
tion on  each  side ) . 

6)  Number  of  scale  rows  between  the 
subocular  scales  and  the  supralabial  scales. 

7)  Number  of  vertical  rows  of  dorsal 
scales  in  a  distance  equal  to  that  from  the 
tip  of  the  snout  to  the  anterior  bony  wall 
of  the  orbit,  this  distance  measured  by  ver- 
nier calipers,  laid  off  on  the  back  about 
three  rows  below  the  median  dorsal  crest 
scales,  and  the  number  of  scales  counted 
under  a  binocular  dissecting  microscope. 

8)  Number  of  horizontal  rows  of  dorsal 
scales  in  the  snout-eye  distance,  the  scales 
counted  at  midbody.  This  count  was  not 
taken  in  most  juveniles  or  on  those  speci- 
mens that  were  shrunken  or  poorly  pre- 
served since,  under  the  latter  circumstances, 
some  smaller  ventrolateral  or  ventral  scales 
will  be  included. 

9)  Number  of  transverse  rows  of  ventral 
scales  in  the  snout-eye  distance. 


10)  Number  of  lamellae  on  phalanges  II 
and  III  of  the  fourth  toe. 

11)  Height  of  the  nuchal  crest  scales, 
categorized  as  very  high,  high,  moderate, 
or  low. 

12)  Height  of  dorsal  crest  scales,  cate- 
gorized as  high,  moderate,  or  low. 

13)  Dorsal  coloration  and  pattern  of 
males  and  females,  separately. 

14)  Ventral  coloration  of  males  and  fe- 
males, separately. 

15)  Color  of  dewlap,  in  males  and  fe- 
males, separately. 

16 )  Color  and  pattern  of  chin  and  throat 
in  males  and  females,  separately. 

17)  Color  and  pattern  of  upper  surface 
of  head  in  males  and  females,  separately. 

18)  Color  of  eyeskin. 

19)  Color  and  pattern  of  upper  surfaces 
of  hindlimbs. 

20)  Color  and  pattern  of  juveniles  and 
subadults. 

The  above  characteristics  are  variously 
useful  as  far  as  delimiting  the  nameworthy 
populations  of  A.  ricordi.  Detailed  com- 
ments on  these  characteristics  are  made  be- 
low, with  especial  attention  to  pitfalls  in 
their  reliability  and  usage. 

1)  The  snout-vent  length  of  mature  in- 
dividuals of  both  sexes  is  remarkably  uni- 
form throughout  the  entire  series.  Males 
are  easily  distinguished  from  females  at 
any  age  by  the  presence  of  one  (occasion- 
ally two)  pairs  of  enlarged  postanal  scales. 
Many  ASFS  specimens  have  the  hemipenes 
extruded.  Males  in  general  reach  a  larger 
snout-vent  length  than  females;  the  largest 
male  (ASFS  V29284)  has  a  snout-vent 
length  of  180  and  is  from  sample  (4). 
whereas  the  largest  female  (ASFS  V31397) 
has  a  snout-vent  length  of  151  and  is  from 
sample  ( 12 ) .  The  mean  difference  in 
snout-vent  lengths  between  the  two  sexes 
is  about  10  mm  in  almost  all  samples  with 
the  exception  of  maximally  sized  individ- 
uals in  both  sexes  in  sample  (2)  where  the 
difference  is  3  mm,  sample  (3)  where  the 
difference  is  27  mm,  sample  (4)  where  the 


HisPANioLAN  Giant  Angle  •  Schwartz 


97 


difference  is  32  mm,  and  sample  (7)  where 
both  sexes  are  of  the  same  size.  Whether 
tliese  exceptions  to  the  10-mm  generahza- 
tion  are  meaningful  is  debatable.  At  least 
samples  (2)  and  (4)  include  Icmg  series 
of  specimens,  and  even  samples  (3)  and 
(7)  include  more  than  ten  individuals. 
Adults  of  the  two  sexes  are  readily  distin- 
guished moi-phologically,  since  males  have 
a  high  tail  "fin"  that  is  supported  by  the 
bony  extensions  of  the  neural  spines;  this 
feature  is  lacking  in  females,  although  they 
may  have  a  much  reduced  caudal  crest  in 
the  form  of  a  low  ridge.  The  terminal  half 
of  the  tails  of  many  males  is  crestless;  I  at 
first  considered  that  this  was  due  to  break- 
age with  subsequent  regeneration  without 
regeneration  of  the  tail  "fin."  Many  speci- 
mens have  this  condition,  however,  without 
any  obvious  change  in  basic  caudal  scale 
shape  and  arrangement,  and  this  is  the  nor- 
mal condition  in  the  tails  of  males.  Often 
the  uncrested  portion  of  the  tail  is  quite 
differently  colored  or  patterned  than  the  re- 
mainder of  the  tail;  such  cases  are  due  to 
regeneration. 

2)  The  number  of  snout  scales  at  the 
level  of  the  second  canthal  has  been  em- 
ployed as  a  characteristic  to  separate  the 
recognized  subspecies.  Williams  ( 1962, 
1965)  recorded  the  following  variation  in 
the  four  subspecies:  ricordi,  7-9;  Ijaleatus, 
2-5;  harahonae,  4-6;  and  leheri,  4-6  (3-6 
on  map,  fig.  2,  1965:  7).  It  should  be  re- 
called that  Williams  himself  pointed  out 
that  this  count  alone  would  not  distinguish 
all  these  taxa  from  each  other.  Certainly 
overlap  between  haleatus  and  ricordi  is 
nonexistent,  and  between  haleatus  on  one 
hand,  and  harahoiiae  or  leheri  on  the  other 
hand,  the  overlap  is  small.  Counts  on  403 
specimens  made  by  myself  do  not  extend 
the  parameters  of  snout  scales  at  all:  within 
the  entire  lot  of  specimens,  these  scales 
vary  from  2  to  9,  just  as  in  Williams's  data. 
However,  the  variation  within  populations 
may  be  much  greater  than  Williams  antici- 
pated. For  instance,  in  sample  (13),  the 
counts  vary  between  2  and  7,  and  in  series 


(12)  between  4  and  8.  Most  samples  have 
three  or  four  categories  of  number  of  snout 
scales.  I  am  in  no  way  implying  that  this 
is  an  invalid  or  poor  character  for  differen- 
tiation of  populations  of  A.  ricordi,  how- 
ever; it  is,  rather,  an  extremely  useful  one 
but  requires  amplification  and  interpreta- 
tion. 

If  we  examine  only  those  samples  (1-8) 
which  are  assigned  to  haleatus,  the  amount 
of  variation  in  snout-scales  is  2-5,  exactly 
that  assigned  to  this  taxon  by  Williams. 
However,  within  the  broad  area  covered  by 
haleatus,  there  are  strong  modalities  of 
snout-scales.  In  samples  (1),  (4),  (5),  and 
(6),  the  mode  is  2  scales,  whereas  in  sam- 
ples (2),  (7)  and  (8),  the  mode  is  4.  Only 
one  sample,  (3),  has  bimodes  of  2  and  4 
scales.  In  harahonae  (9)  the  range  is  2- 
5  (mode  4).  In  those  samples  which  are 
associated  with  nominate  ricordii  (samples 
11  and  12),  leheri  (sample  13)  and  adja- 
cent Castillon  material  (sample  14),  the 
range  is  2-9,  thereby  showing  complete 
overlap  in  range  of  this  count  with  that  of 
both  haleatus  and  harahonae.  In  fact,  in 
leheri  ( 13 )  alone,  the  range  of  snout  scales 
(2-7)  almost  embraces  that  for  all  other 
samples  and  thus  the  entire  species.  Mo- 
dalities in  this  complex  of  samples  are  5 
(sample  11),  7  (sample  12),  4  (sample  13) 
and  6  (sample  14).  Sample  (7)  is  nomi- 
nate ricordi. 

3 )  The  number  of  vertical  rows  of  loreal 
scales  ranges  from  5  to  10.  The  greatest 
variability  is  in  samples  (1),  (2),  (7),  and 
(12),  where  the  row  counts  in  each  case 
are  5-9,  6-10,  5-9,  and  6-10.  Most  samples 
have  four  categories  of  number  of  loreal 
rows.  The  modes  vary  as  follows:  5  (sam- 
ple 11),  6  (samples  8,  9,  13),  7  (samples  1, 
3,  4,  5,  6,  7,  12,  and  14),  and  8  (.sample  2). 

4)  The  minimal  number  of  scales  be- 
tween the  supraorbital  semicircles  varies 
between  1  and  5;  no  specimen  has  the  semi- 
circles in  contact.  Modes  in  general  are 
very  strong,  and  the  usual  mode  is  3  scales 
(samples  1-7;  sample  8  has  a  bimode  of  2 
and  3  scales);  these  are  all  samples  that 


\ 


98 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


are  assigned  to  baleatiis.  A  mode  of  3 
scales  occurs  also  in  samples  (11)  and 
( 13 ) ,  and  of  4  in  samples  ( 12 )  and  ( 14 ) . 

5)  The  number  of  scales  between  the  in- 
terparietal scale  and  the  supraorbital  semi- 
circles varies  between  3/3  and  7/7.  Modes 
(which  in  some  cases  are  quite  strong  and 
in  others  less  so)  are:  4/4  (samples  1,  5,  6, 
8,  9,  11,  13,  14)  and  5/5  (samples  2,  4,  7, 
12).  Sample  (3)  is  peculiar  in  having  the 
mode  4/5  (six  of  16  lizards),  with  adjacent 
counts  of  4/4  (four  lizards)  and  5/5  (five 
lizards).  There  is  high  variability  in  .this 
count;  it  can  be  assessed  in  another  fashion, 
namely,  the  frequency  with  which  any 
scale  count  (i.e.,  4,  5,  6,  etc.)  occurs  within 
the  sample,  regardless  of  its  pairing  with 
another  count  on  the  other  side  of  the  head. 
With  the  use  of  this  technique,  the  fre- 
quency of  involvement  of  4  scales  in  the 
inteiparietal-semicircle  contact  varies  be- 
tween 43  percent  (sample  1)  and  67  per- 
cent (sample  8),  and  of  5  scales  between 
52  percent  (sample  2)  and  64  percent 
( sample  7 ) .  Of  the  entire  lot  of  specimens, 
there  is  only  one  occurrence  of  7/7  in  this 
position  ( sample  12 )  but  many  occurrences 
of  3/3  (samples  1,  5,  6,  9,  11,  12,  13,  a  total 
of  12  incidences). 

6)  The  number  of  scale  rows  between 
the  subocular  scales  and  the  supralabials  is 
fairly  constantly  1,  and  this  is  a  strong 
mode  or  the  exclusive  category  in  all  sam- 
ples except  sample  ( 13 ) .  In  this  lot  ( which 
is  the  type  series,  with  a  few  new  speci- 
mens, of  leberi),  48  percent  of  the  lizards 
have  the  suboculars  and  supralabials  in 
contact.  Elsewhere,  contact  is  absent  in 
samples  (7)  and  (11).  The  frequency  of 
contact  varies  in  all  other  samples  between 
3  percent  (sample  9)  and  17  percent  (sam- 
ples 6,  8,  and  14).  These  three  samples  are 
widely  separated  geographically  and  the 
frequency  in  none  of  them  even  approaches 
the  very  high  incidence  of  contact  in  sam- 
ple ( 13 ) .  However,  it  is  noteworthy  that 
samples  (13)  and  (14)  are  adjacent  geo- 
graphically. 

7),  8),  9)  In  reference  to  all  counts  in- 
volving laying  out  the  snout-orbit  distance 


on  the  body,  I  can  do  no  better  than  once 
more  to  reiterate  the  cautions  previously 
expressed  ( Schwartz,  1964;  Garrido  and 
Schwartz,  1968;  Schwartz  and  Garrido, 
1972)  in  reference  to  Anolis  equestris  and 
members  of  the  genus  Chamaeleolis.  For 
these  standard-distance  counts,  old  and 
poorly  preserved,  limp,  curled,  uninjected, 
or  otherwise  distorted  specimens  are  much 
less  useful  and  reliable  than  properly  pre- 
served, injected,  and  positioned  lizards. 
Luckily,  by  far  the  largest  quantity  of  A. 
ricordi  under  study  are  well  preserved. 
However,  I  have  abandoned  counts  of  hor- 
izontal dorsals  on  young  juveniles,  even 
well-preserved  ones,  or  on  any  adults 
whose  condition  precluded  taking  these 
counts  meaningfully.  The  juvenile  situation 
is  peculiar  in  that  invariably,  despite  the 
relatively  shorter  snout  of  young  specimens, 
laying  out  this  distance  to  count  horizontal 
rows  involved  including  several  rows  of  ex- 
tremely lateral  and  ventral  scales,  which 
are  smaller  than  true  dorsals  and  which 
thus  tend  to  increase  the  horizontal  counts. 
I  have  taken  vertical  dorsals  and  ventrals 
on  juveniles,  however,  and  they  do  not 
skew  the  data.  Of  the  three  standard-dis- 
tance counts,  those  of  vertical  dorsals  and 
ventrals  are  much  more  reliable  than  are 
those  of  horizontal  dorsals. 

Vertical  dorsal  scale  rows  vary  between 
12  and  26,  with  the  low  count  in  sample 
( 4 )  and  the  high  count  in  sample  ( 12 ) . 
Means  range  from  15.4  (sample  4)  to  21.1 
(sample  12).  These  two  represent,  respec- 
tively, lots  assignable  to  baleatus  and  ri- 
cordi, sensu  lato.  No  generalizations  of 
mean  number  of  vertical  dorsals  in  refer- 
ence to  samples  and  geography  can  be 
made,  since  the  range  in  samples  now  as- 
sociated with  baleatus  varies  between  15.4 
and  19.2,  with  ricordi  17.3  and  21.1,  bara- 
honae  17.2,  and  leberi  16.5  and  16.7.  Com- 
parisons and  significance  of  vertical  dorsal 
scale  counts  are  shown  in  Table  I. 

Number  of  horizontal  dorsal  rows  ranges 
from  13  (sample  1)  to  34  (sample  7). 
Means  vary  between  17.1  (sample  11)  to 
24.6  (sample  12).  The  latter  sample  is  that 


HisPANioLAN  Giant  Angle  •  Schwartz 


99 


Table  I. 


Taxon 


Comparison  of  number  of  vertical  dorsal  scales  in  popxjtlations 
OF  giant  Hispaniolan  angles 


M    (±2 

standard 

error   of 

mean) 


.A 


50 

c 
a 


c 
c 


s 

2 

a. 

■2 

"a 
£ 

1 

-*- 

J 

c 

+ 

+ 

+ 

+ 

+ 

+ 

+ 

— 

+ 

— 

+ 

— 

+ 

— 

+ 

— 

+ 

+ 

+ 

— 

+ 

+ 

+ 

— 

— 

+ 

+ 

+ 

+ 

+ 

+ 

ricordi  50 

leheri  54 

stihsolanus  15 

harahonae  33 

samanae  54 

caendcolatus  37 

litorisdva  16 

scelestus  60 

multistrtippus  20 

sublimis  18 

baleatus  15 


21.1  ± 

16.5  ± 

17.3  ± 

17.2  ± 

16.6  ± 

17.1  ± 
15.9  ± 

15.4  ± 
18.6  ± 

19.2  ± 


.57 
.46 
.96 
.67 
.46 
.72 
.75 
.45 
1.06 
.70 


+ 


+        + 


+ 


+ 


+ 


17.5  ±  1.16 


A.  r.  victilus,  A.  b.  alboceUatus,  and  A.  b.  fraiidator  are  not  included.  A  "-)-"  in  the  table  indicates  that  the  two 
subspecies  involved  differ  significantly  ( non-overlap  of  two  standard  errors  of  mean ) ;  a  "— "  indicates  no  statistical 
difference.  Note  that  the  mean  of  A.  r.  ricordi  differs  significantly  from  the  means  of  all  other  taxa;  that  of  scelestus 
differs  significantly  from  those  of  all  other  taxa  except  litorisilva;  and  that  of  sublimis  differs  significantly  from  those 
of   all   other  taxa   except   multistruppus   and   baleatus. 


of  nominate  ricordi,  the  former  a  peripheral 
isolate  of  baleatus. 

Ninnber  of  transverse  rows  of  vential 
scales  varies  between  15  (samples  1  and 
13 )  and  34  ( sample  7 ) .  Means  range  from 
20.2  (sample  13)  to  2.5.1  (sample  6);  sam- 
ple ( 13 )  is  leheri. 

10)  The  number  of  lamellae  on  pha- 
langes II  and  III  of  the  fourth  toe  varies 
between  27  and  39.  The  variation  in  any 
population  is  so  great  that  this  count  is 
meaningless  as  far  as  differentiation  be- 
tween any  populations  of  A.  ricordi. 

11),  12)  Williams  (1962,  1965)  de- 
scribed the  relative  heights  and  lengths  of 
both  the  nuchal  and  dorsal  body  crest 
scales.  Certainly  the  differences  between 
baleatus  and  ricordi,  for  instance,  are  so 
very  obvious  on  casual  examination  that 
one  has  no  difficulty  in  ascertaining  the 
taxon  involved.  Williams  also  pointed  out 
(1962:  4-5)  that  in  some  specimens  there 
is  "sometimes  a  regular  alternation  of  rela- 
tively high  triangular  single  scales  and 
pairs  of  much  lower,  more  quadrangular 
scales,"  with  the  result,  on  some  specimens, 
of  double  crest  scale  rows  on  the  neck;  the 
net  effect  of  this  condition  is  a  rather  in- 
discriminate grouping  or  elongate  patch  of 


multiple  nuchal  crest  scales.  Although  tliis 
condition  occurs  erraticallv,  it  seems  to  be 
most  predominant  in  specimens  from  the 
Tiburon  Peninsula,  but  it  occurs  elsewhere 
in  nominate  ricordi  and  even  occasionally 
in  specimens  assigned  now  to  baleatus. 
Such  a  "hypertrophied"  nuchal  crest  condi- 
tion does  not  completely  fit  any  logical  geo- 
graphical pattern  nor  is  it  totally  consistent 
within  any  sample,  although  there  are  ten- 
dencies toward  it  as  noted  above.  In  any 
event,  it  does  not  obscure  the  height  of  the 
nuchal  crest  scales. 

In  my  own  analysis,  I  have  used  a 
slightly  different  method  in  recording 
height  of  crest  scales.  Nuchal  scales  were 
recorded  for  each  specimen  as  very  high, 
high,  moderate,  or  low.  Such  a  verbal  quan- 
tification is  not  totally  satisfactory,  since 
the  investigator's  impressions  may  change 
as  the  study  progresses.  To  avoid  this  pit- 
fall, I  re-examined  many  specimens  that 
had  been  studied  earlier  in  the  work  and 
reconfirmed  my  own  early  impressions  with 
my  later  ones.  Body  crest  scales  were  re- 
corded as  high,  moderate,  and  low. 

In  nuchal  crest  scales,  very  high  scales 
are  those  which  are  very  elongate,  attenu- 
ate, almost  spinelike  ( but  of  course  flexible, 


100 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


not  stiff),  with  the  base  much  shorter  than 
the  height  of  the  scale.  High  scales  are 
those  which  are  shorter  and  less  attenuate 
than  very  high  scales,  but  whose  height  is 
still  much  greater  than  the  base.  Moderate 
scales  are  lower  and  not  attenuate,  although 
they  may  be  pointed,  with  the  height  and 
base  about  equal  in  length.  Low  scales  are 
lower  than  long.  The  same  categories  and 
interpretations  apply  to  body  crest  scales, 
although  no  lizard  has  the  body  crest  scales 
so  high  as  the  nuchal  scales. 

Several  other  points  are  pertinent.  I  have 
not  used  this  datiun  from  juvenile  and  sub- 
adult  lizards  ( all  lizards  below  100  mm 
snout-vent  length)  since  it  is  obvious  that, 
regardless  of  the  taxon  or  sample,  all  young 
A.  ricordi  have  low  nuchal  and  dorsal  crest 
scales,  which,  as  the  lizard  matures,  be- 
come increasingly  more  specialized  until 
the  adult  condition  is  reached  at  about  110 
mm  snout-vent  length.  Thus  young  ricordi 
and  young  baleatus,  two  taxa  that  are  re- 
markably distinct  in  this  feature  as  adults, 
are  identical  in  crest  development. 

Secondly,  it  might  be  assumed  that  (es- 
pecially) nuchal  crest  scales  might  be  bet- 
ter developed  in  adult  males  than  in  adult 
females;  this  excessive  development  might 
reasonably  be  assumed  or  construed  as  a 
sexually  dimorphic  character,  with  hyper- 
development  in  males.  Such  does  not  seem 
to  be  the  case,  and  many  female  baleatus, 
for  instance,  have  very  high  nuchal  crest 
scales,  as  high  as  those  of  males.  In  fact, 
comparisons  of  males  and  females  of  indi- 
vidual samples  show  that,  within  each  sam- 
ple, there  is  remarkable  consistency  be- 
tween frequencies  of  the  very  high,  high, 
and  moderate  categories  in  both  sexes. 

Thirdly,  as  one  might  expect,  there  is  a 
sequence  of  crest  scale  heights  in  the  nu- 
chal-body series.  If  the  nuchal  scales  are 
very  high,  the  body  scales  are  high;  if  the 
nuchal  scales  are  moderate,  the  body  scales 
are  low,  etc.  In  no  case  have  I  recorded  a 
transition  from  very  high  nuchal  crest 
scales  to  moderate  body  scales,  for  exam- 
ple. There  is  thus  a  definite  correlation  be- 
tween height  of  nuchal  scales  and  those  of 
the  body  crest. 


13) -19)  The  color  and  pattern  details  of 
Anolis  ricordi  throughout  its  range  need 
not  be  gone  into  in  detail  at  this  point.  It  is 
now  sufficient  to  acknowledge  that  these 
lizards  show  metachrosis  varying  from 
shades  of  green  and  green-gray  to  brown. 
The  pattern  elements,  however,  are  quite 
stable,  although  the  hues  involved  in  the 
pattern  itself  may  change  with  changing 
base  colors.  There  is  little  evidence  to  in- 
dicate that  a  lizard  which  is,  for  instance, 
blotched  in  one  color  phase  will  become 
crossbanded  in  another.  Thus,  despite 
changes  in  hues  and  even  in  base  colors, 
patterns  remain  constant.  It  is  of  interest 
to  note  that  greens  seem  to  be  the  colors 
that  predominate  in  the  wild.  The  few  A. 
ricordi  that  I  have  seen  during  the  day 
have  always  been  green.  The  many  lizards 
that  I  have  seen  and  collected  at  night 
were  almost  always  green,  and  usually  at 
their  maximum  of  pattern  expression  while 
they  slept.  It  is  this  nocturnal  assumption 
of  the  green  phase  coloration  that  renders 
these  lizards  so  very  conspicuous  at  night 
while  they  sleep  on  exposed  branches, 
vines,  or  among  the  arboreal  greenery. 
Even  in  those  populations  ( Boca  de  Yuma, 
Sierra  de  Baoruco)  in  which  the  greens  in- 
volved are  not  bright,  the  lizards  are  still 
quite  obvious  at  night.  It  is  only  rarely 
that  one  encounters  a  brown  A.  ricordi  at 
night.  I  have  notes  on  only  one  instance  of 
this  condition,  a  subadult  lizard  from  the 
Cordillera  Septentrional. 

15)  The  dewlap  coloration  in  A.  ricordi 
is  variable.  In  some  populations,  males 
have  a  pale  yellow  to  gray  dewlap,  whereas 
in  others  the  males  have  dewlaps  that  are 
peach  or  vivid  orange.  It  should  be  noted 
that  both  sexes  in  A.  ricordi  have  dewlaps 
and  that  the  general  hue  of  the  female 
dewlap  is  like  that  of  the  males,  except  that 
basally  it  is  usually  streaked  with  browns, 
dark  grays,  or  charcoal,  and  the  ground 
color  is  less  bright  than  that  of  males.  In 
some  regions,  the  female  dewlap  is  quite 
differently  colored  than  that  of  males. 

20)  Juveniles  and  subadults  present  sev- 
eral problems  that  are  presently  insoluble. 
I  suspect  that  much  will  be  revealed  once 


HisPANioLAN  Giant  Angle  •  Schwartz 


101 


we  know  the  repertory  of  pattern  and  color 
in  yoinig  individuals,  but  these  data  are 
not  axailable  nx)w.  Although  there  are 
many  young  specimens  at  hand,  they  are 
inconsistent  within  populations  or  even 
small  samples.  The  juxenile  color  is  nor- 
mally some  shade  of  green  (or  browns  un- 
der stress),  most  often  with  two  to  four 
pale  (cream,  whitish,  pale  gray)  cross- 
bands.  Many  small  specimens  are  a  uni- 
form green  without  any  dorsal  markings. 
In  other  juveniles,  the  dorsum  has  many 
conspicuous  crossbands  with  two  shades  of 
greens  (or  browns),  separated  by  promi- 
nent bands  of  pale  greens  (yellow-green, 
pea-green),  to  give  a  very  tigroid  lizard; 
this  condition  persists  into  the  adults  of  one 
population,  as  does  the  more  simply 
banded  juvenile  pattern  noted  above  in 
other  populations.  The  juveniles  usually 
have  dark  gray  to  charcoal  dewlaps,  re- 
gardless of  their  sexes,  and  often  there  are 
charcoal  or  white  markings  on  the  neck  or 
aboN'c  the  forelimb  insertion  or  somewhere 
anteriorly.  These  variants  are  discussed  un- 
der each  subspecies  below,  and  there  is  no 
need  to  go  into  the  details  here.  However, 
I  do  wish  to  point  out  that  ju\'enile  patterns 
are  more  variable  than  are  those  of  adults, 
and  that  I  do  not  know  how  to  interpret 
this  situation. 

SYSTEMATIC  ACCOUNTS 

AnoWs  ricordi  Dumeril  and  Bibron 

Anolis   ricordii   Dumeril   and   Bibron,    1837.     Erp. 
gen.,  4:  167. 

Type  locality.  St.-Domingue;  holotype. 
Museum  National  d'Histoire  Natmelle  1272. 

Definition.  A  giant  species  of  Hispan- 
iolan  Anolis  characterized  by  the  combina- 
tion of  moderate  size  (males  to  160  mm, 
females  to  151  mm  snout-vent  length), 
snout  scales  at  level  of  second  canthal 
scales  variable,  between  2  and  9  (modally 
4,  5,  6,  or  7  by  population)  but  usually  4 
to  9  (97  percent),  vertical  loreal  rows  5  to 
10  (modes  by  population  5,  6,  and  7), 
scales  between  supraorbital  semicircles  2  to 
5  (modes  3  or  4  by  population),  inteipari- 
etal  scale  separated  from  supraorbital  semi- 
circles modally  by  4  or  5  scales,  vertical 


dorsal  scale's  generally  small  ( 14  to  26  in 
standard-distance),  ventral  scales  relatively 
large  (15  to  32  in  standard-distance),  nu- 
chal crest  scales  in  both  sexes  moderate  to 
low,  never  \'ery  high  or  high,  body  crest 
scales  usually  low,  subocular  scales  usually 
not  in  contact  with  supralabials  but  one 
population  is  remarkably  exceptional  in 
this  character;  dorsal  body  coloration  and 
pattern  some  shade  of  green,  in  some  geo- 
graphic regions  flecked  irregularly  with 
paler  and  darker  green  to  give  a  beadwork 
effect;  male  body  pattern  either  of  irregular 
black  to  dark  brown  blotches  on  the  neck, 
occipital  region  of  the  head,  and  on  sides 
(often  delimiting  two  pale  longitudinal 
bands)  or  with  three  longitudinal  dark 
brown  stripes  on  each  side  or  with  dark 
saddles  and  a  bluish  green  flank  stripe  or 
with  a  powdery  pale  blue-green  lateral 
stripe;  females  usually  without  dark  dorsal 
or  lateral  markings  although  in  some  areas 
females  have  a  darker  brown  reticulum, 
three  pale  gray  to  yellow  narrow  cross- 
bands,  longitudinal  black  lines,  or  two  pur- 
ple flank  stripes;  dewlap  in  males  variable, 
from  yellowish  gray  to  peach,  bright  or- 
ange, or  deep  yellow,  in  females  from 
peach  to  deep  yellow  or  dull  orange  or 
even  inky  brown  or  inky  blue-black,  chin 
and  throat  dull  yellow,  yellow-green,  or 
pale  blue-green  in  males,  eyeskin  dark 
( charcoal,  dark  brown )  to  light  ( pale  blue ) 
in  males,  charcoal  to  pale  green  in  females, 
and  usually  with  a  prominent  pale  subocu- 
lar semicircle  clearly  delineated. 

Distribution.  The  Tiburon  Peninsula  in 
Haiti,  east  to  the  vicinity  of  Saltron,  Dept. 
de  rOuest,  thence  north  to  the  northern 
Haitian  coast  from  Port-de-Paix  east  to  Ter- 
rier Rouge  and  thence  into  the  Republica 
Dominicana  east  as  far  as  the  vicinity  of 
Los  Quemados,  Santiago  Rodriguez  Prov- 
ince, south  to  Restauracion,  Dajabon  Prov- 
ince; also  extending  from  Haiti  onto  the 
southwestern  slopes  of  the  Cordillera  Cen- 
tral in  San  Juan  Province  and  in  the  Sierra 
de  Neiba  in  La  Estrelleta  and  San  Juan 
provinces;  altitudinal  distribution  from  sea 
level  to  elevations  of  about  4000  feet  ( 1220 
meters)  south  of  Castillon,  Dept.  du  Sud, 


102         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


Haiti,  and  to  3500  feet  ( 1068  meters )  west 
of  Marmelade  in  the  Chaine  de  Marmelade, 
Dept.  de  I'Artibonite,  and  3400  feet  ( 1037 
meters)  south  of  Elias  Pifia  in  the  Sierra  de 
Neiba,  La  Esti-elleta  Province,  RepubHca 
Dominicana. 

Anofis  ricordi  ricordi  Dumeril  and  Bibron 

Type  locality.  "St.-Domingue";  here  re- 
stricted to  the  vicinity  of  Port-au-Prince, 
Dept.  de  I'Ouest,  Haiti. 

Definition.  A  subspecies  of  A.  ricordi 
characterized  by  the  combination  of  mod- 
ally  7  snout  scales  between  second  canthal 
scales,  7  vertical  rows  of  loreal  scales,  4 
scales  between  the  supraorbital  semicircles, 
5/5  scales  between  the  interparietal  and  the 
supraorbital  semicircles,  high  number  of 
vertical  dorsal  scales  (17-26;  mean  21.1), 
high  number  of  ventral  scales  ( 19-32;  mean 
24.7 ) ,  nuchal  crest  scales  moderate  ( rarely ) 
to  low  (usually),  body  crest  scales  moder- 
ate (rarely)  to  low  (usually),  subocular 
scales  usually  in  contact  with  supralabial 
scales;  males  usually  with  some  black  lat- 
eral markings  on  the  neck  and  thoracic  re- 
gion, and  on  the  occipital  region  of  the 
head,  but  at  times  these  markings  are  ab- 
sent (see  discussion  below),  females  green 
and  without  definite  black  lateral  markings 
but  at  times  reticulate  with  brown,  the  re- 
ticulum delimiting  a  pair  of  clear  green  lat- 
eral stripes  or  with  three  pale  gray  to  green 
vertical  narrow  bars;  dewlap  variable,  in 
males  from  peach  or  pale  peach  to  gray  or 
yellowish  gray,  and  in  females  from  peach 
to  blue-gray  or  inky  blue  or  inky  black  ( see 
discussion  beyond). 

Distribution.  Northern  Haiti  from  Port- 
de-Paix  east  to  Terrier  Rouge  and  into  the 
Republica  Dominicana  as  far  east  as  the 
vicinity  of  Los  Quemados,  Santiago  Ro- 
driguez Province,  and  as  far  south  as  Re- 
stauracion,  Dajabon  Province,  south  in  Haiti 
to  the  Port-au-Prince  region  ( Morne  De- 
cayette,  Diquini,  Petionville),  as  well  as 
east  into  the  Republica  Dominicana  in  the 
Sierra  de  Neiba  and  the  southwestern 
slopes  of  the  Cordillera  Central  in  La  Es- 
trelleta  and  San  Juan  provinces. 


Discussion.  I  have  little  doubt  that  the 
extensive  range  that  I  here  ascribe  to  nom- 
inate A.  ricordi  is  incorrect.  There  are  sev- 
eral very  obvious  differences  in  coloration 
and  pattern  between  northern  and  south- 
ern specimens  of  A.  r.  ricordi;  thus  the  defi- 
nition of  the  subspecies,  in  order  to  include 
all  pattern  variants,  is  necessarily  cumber- 
some. The  problem  is  presently  unresolv- 
able  since,  other  than  the  series  from  near 
Port-au-Prince  and  the  specimens  from 
northern  Haiti,  there  are  huge  areas  in 
Haiti  whence  specimens  remain  unknown. 
A  detailed  discussion  of  the  chromatic  and 
pattern  features  in  the  various  segments  of 
A.  r.  ricordi  is  given  below. 

The  series  of  50  specimens  assigned  to 
the  nominate  subspecies  shows  the  follow- 
ing variation.  The  largest  male  (ASFS 
V31395)  has  a  snout-vent  length  of  160, 
the  largest  female  (ASFS  V31397)  151; 
both  are  from  4.1  mi.  NW  Juan  de  Herrera, 
San  Juan  Province,  Republica  Dominicana. 
Snout  scales  at  level  of  second  canthals 
vary  between  4  and  8;  the  mode  is  7  (22 
specimens).  The  vertical  loreal  rows  vary 
between  6  and  10,  with  a  mode  of  7  (20 
specimens).  There  are  between  3  and  5 
scales  between  the  supraorbital  semicircles 
(mode  4).  There  are  modally  5  scales  be- 
tween the  interparietal  and  the  semicircles; 
5  scales  are  involved  in  53  percent  of  the 
combinations;  actual  counts  are  3/3  (1), 
4/4  (10),  4/5  (8),  5/5  (17),  5/6  (9),  6/6 
(2),  7/7  (1),  and  5/7(1).  Vertical  dorsals 
range  between  17  and  26  (mean  21.1),  hor- 
izontal dorsals  between  19  and  30  (24.6), 
and  ventrals  between  19  and  32  (24.7).  Of 
28  adult  males,  six  have  moderate  nuchal 
crest  scales  and  22  have  these  scales  low; 
of  11  females,  all  have  the  nuchal  crest 
scales  low.  Body  crest  scales  are  moderate 
in  two  males  and  low  in  26,  and  11  females 
have  the  body  crest  scales  low.  The  sub- 
oculars  are  separated  from  the  supralabials 
in  45  of  49  instances,  and  contact  between 
these  scales  occurs  in  four  lizards  (8  per- 
cent ) . 

The  southern  specimens  from  the  Port- 
au-Prince  region  and  including  two  from 


HisPANioLAN  Giant  Anole  •  Schwart::. 


103 


Mirebalais,  consist  of  ten  adult  males,  three 
adult  females,  and  two  juvenile  females 
(MCZ  60013-14).  The  latter  two  speci- 
mens (with  snout-vent  lengths  of  89  and 
92  mm)  can  be  easily  dismissed  in  that 
they  are  presently  patternless  green.  Color 
notes  in  life  on  one  southern  male  (ASFS 
V9024)  state  that  in  the  green  phase,  the 
lizard  had  the  dorsum  a  mixture  of  pale 
green,  brown,  and  yellow,  with  green  the 
predominant  color,  the  net  effect  being  one 
of  bead  work.  The  upper  surface  of  the 
head  was  a  mixture  of  pinkish  and  yellow 
scales,  the  mental  region  and  adjacent  up- 
per labials  were  dull  yellow,  and  the  venter 
pale  green.  The  dewlap  was  peach  with 
the  dewlap  scales  yellow.  All  males  (with 
the  exception  of  MCZ  69404,  which  is  an 
albino )  have  some  black  to  dark  brown  oc- 
cipital blotching,  usually  extending  onto 
the  neck  and  thence  onto  the  area  above 
the  forelimb  insertion.  The  extent  of  the 
dark  anterior  markings  is  variable,  but  they 
are  present  in  all  males  and  quite  vivid  in 
freshly  taken  specimens.  A  pale  subocular 
crescent  is  present,  often  extending  posteri- 
orly to  form  a  pale  preauricular  blotch, 
bounded  above  by  a  large  dark  temporal 
blotch  that  may  form  an  occipital  chevron 
by  joining  its  mate  on  the  other  side. 
Southern  females  were  recorded  in  life  as 
pale  to  bright  green  without  any  dark  pat- 
tern, with  a  moderately  well-defined  pale 
subocular  crescent  that  may  expand  into  a 
pale  preauricular  blotch;  the  venters  were 
yellow-green.  In  one  female,  the  dewlap 
was  reported  as  blue-gray  with  yellow- 
green  streaks.  The  hindlimbs  are  not  prom- 
inently banded.  Neither  sex  has  the  throat 
marked  with  any  blotching  or  dotting,  al- 
though the  females  have  the  throat  some- 
what clouded  with  dull  dark  green. 

The  series  for  the  Sierra  de  Neiba  and 
the  Cordillera  Central  consists  of  six  males, 
four  females,  and  eight  juveniles  and  sub- 
adults.  The  males  were  described  in  life 
as  being  dark  green  with  pale  green  cross- 
bands,  or  pale  green  with  four  paler  green 
crossbands,  or  patternless  green.  The  up- 
per surface  of  the  head  was  creamy  tan. 


black  blotches  occur  in  this  series  of  males 
also,  but  the  occiput  lacks  clearly  defined 
black  areas,  and  the  black  on  the  body  is 
much  more  extensive  than  it  is  in  all  south- 
ern males,  the  extreme  condition  being  that 
shown  by  ASFS  V31395,  which  has  exten- 
sive black  blotching  over  two-thirds  of  the 
back  and  sides.  The  pale  subocular  cres- 
cent is  very  obscure,  but  there  is  a  promi- 
nent pale  preauricular  spot  in  most  males. 
The  females  from  this  region  are  plain 
green,  without  dark  markings,  and  there  is 
a  prominent  pale  supralabial  blotch  in  the 
area  which  in  males  is  occupied  by  the  pale 
preauricular  blotch.  Dewlaps  in  males  and 
females  were  invariably  recorded  as  peach, 
and  both  sexes  had  charcoal  eyeskins.  As 
in  southern  specimens,  the  chins  and 
throats  are  pale  green  and  without  any  defi- 
nite markings,  except  that  the  throats  of  fe- 
males are  sufi^used  with  darker  green.  The 
eight  juveniles  and  subadults  range  in 
snout-vent  length  from  68  mm  to  92  mm. 
The  two  smallest  specimens,  a  male  and  a 
female,  were  rich  pea-green  in  life  with 
four,  narrow,  cream  transverse  crossbars, 
and  the  smaller  had  in  addition  black 
streaking  in  the  green  areas  and  a  black 
postauricular  smudge.  The  ventral  color 
was  rich  pea-green  and  the  dewlap  skin 
was  blue-black.  All  juveniles  and  subadults 
with  snout-vent  lengths  of  between  76  and 
92  were  bright  emerald-green  dorsally  and 
without  any  dorsal  dark  or  pale  markings; 
one  female  juvenile  ( snout-vent  length  78 ) 
had  a  lateral  black  nuchal  spot  followed  by 
a  bright  yellow  preaxillary  bar,  as  well  as 
a  bright  yellow  subocular  mark.  The  dew- 
lap was  recorded  as  black  in  a  juvenile  fe- 
male with  a  snout-vent  length  of  89.  Of 
the  subadults,  the  most  peculiar  is  a  male 
(ASFS  V31323)  with  a  snout-vent  length 
of  90  which  shows,  as  preserved,  a  vague 
series  of  vertical  lateral  pale  and  dark 
areas,  but  as  yet  no  black  blotching  typical 
of  adult  males. 

The  northern  specimens  are  six  males  and 
one  female  from  Haiti,  five  males  and  two 
females  from  the  Republica  Dominicana, 
and  one  Haitian  subadult  and  two  Domin- 


104         Bulletin  Museum  of  Comparative  Zoology,  \o\.   146,  No.  2 


ican  subadult  and  juvenile  lizards.  Haitian 
males  are  not  only  quite  different  from 
northern  Dominican  males,  but  they  are 
also  strikingly  different  from  central  and 
southern  males.  In  the  Haitian  males,  the 
dorsum  is  gray-green  with  yellow-green 
flecking,  or  a  beadwork  mixture  of  dark  and 
light  green  scales.  The  upper  surface  of  the 
head  is  dark  with  light  flecking,  and  in  one 
male  the  head  was  recorded  as  dark  brown 
with  the  centers  of  the  scales  pale  purple. 
No  male  has  any  occipital  dark  blotching, 
and  any  body  blotching,  if  it  is  at  all  pres- 
ent, is  extremely  restricted  and  maximally 
expressed  as  small  black  areas  above  the 
forelimb  insertion  ( MCZ  66147).  The  pale 
subocular  crescent  is  obscure,  but  there  is 
a  pale  postlabial  line  leading  to  the  auricu- 
lar opening.  Northern  Dominican  males, 
on  the  other  hand,  are  brightly  colored  and 
have  extensive  black  neck  and  side  mark- 
ings; 'n  two  males  these  latter  extend  far 
posteriorly  on  the  body  and  tend  to  delimit 
two  lateral  stripes  on  each  side.  The  upper 
surfaces  of  the  head  are  not  mottled  but 
are  pale  uniform  tan.  In  life,  the  pale  sub- 
ocular  crescent  is  bold  and  pale  blue  to 
white,  and  it  may  extend  to  the  auricular 
opening.  In  Haitian  males,  the  dewlap  is 
grayish  to  yellowish  peach  (pi.  12C5;  all 
color  designations  from  Maerz  and  Paul, 
1950),  pale  gray-green  (about  pi.  19B2), 
or  yellowish  gray  (about  pi.  20B1).  In 
northern  Dominican  males,  the  dewlap  is 
pale  peach  to  pale  yellow  or  grayish  yellow, 
and  the  dewlap  may  be  speckled  with 
brown  basally. 

The  single  Haitian  female  is  presently 
unmarked  green,  with  faint  scattered  cream 
flecking.  The  larger  of  the  two  Dominican 
females  was  pale  green  above  with  a  darker 
brown  reticulum  outlining  a  pair  of  green 
lateral  stripes  on  each  side.  There  was  a 
postauricular  brown  smudge,  followed  by  a 
pale  blue  axillary  smudge.  The  temples 
were  yellow-green,  the  lores  pale  blue  and 
brown,  the  eyeskin  pale  green,  and  there 
was  a  pale  blue  subocular  crescent  that  ex- 
tended into  a  preauricular  pale  blotch.  The 
top  of  the  head  was  marbled  pale  tan  and 


dark  brown,  and  the  venter  was  the  same 
color  as  the  dorsum.  The  other  Dominican 
female  was  green  without  any  dorsal  mark- 
ings. 

The  Haitian  subadult  (MCZ  66148)  is  a 
female  with  a  snout-vent  length  of  106.  It 
is  speckled  with  pale  scales  on  a  dark 
ground  like  Haitian  males.  The  smaller  of 
the  two  Dominican  males  (ASFS  V18008) 
has  a  snout-\'ent  length  of  75  and  was 
bright  yellow-green  above  with  two  cream 
crossbands  and  a  yellow  subocular  cres- 
cent. The  second  Dominican  male  (ASFS 
V32160)  has  a  snout-vent  length  of  103 
mm,  and,  like  Dominican  adult  males,  has 
extensive  black  blotching  on  the  head, 
neck,  and  almost  the  entire  dorsum.  The 
ground  color  was  pale  green,  and  the  dew- 
lap was  dark  brown. 

To  summarize  all  the  above  data,  it  is 
obvious  that  I  have  included  several  popu- 
lations in  A.  r.  ricordi  which  differ  rather 
strikingly  among  themselves.  Southern 
Haitian  males  are  marked  with  black  on 
the  occiput,  neck,  and  anterior  sides,  and 
central  Dominican  and  northern  Domini- 
can specimens  increase  this  tendency  to 
show  even  more  extensive  black  lateral 
markings.  On  the  other  hand,  northern 
Haitian  males  as  a  group  show  very  little 
or  no  black  markings  and  are  basically 
green-flecked  green  lizards.  Northern  Do- 
minican males  are  much  more  colorful  than 
specimens  from  elsewhere,  and  much  more 
contrastingly  marked.  On  the  other  hand, 
all  females  are  fairly  similar,  with  the  ex- 
ception of  the  remarkably  colored  and  pat- 
terned female  from  the  northern  Republica 
Dominicana.  I  suspect  that  it  will  ulti- 
mately be  shown  that  there  are  at  least  two 
more  nameworthy  populations  included  in 
A.  r.  ricordi  as  here  defined  by  me:  a  north- 
ern Haitian  subspecies,  a  northern  and  cen- 
tral Dominican  population,  as  well  as  the 
southern  Haitian  one.  But  the  specimens 
at  this  time  are  from  such  disjunct  localities 
and  are  so  limited  in  number  that  I  am  un- 
willing to  make  the  suggested  nomencla- 
tural  additions. 

Remarks.     A.  r.  ricordi  occurs  in  a  wide 


HisPANioLAN  Giant  Anole  •  Schwartz 


105 


variety  of  situations  but  is  of  course  always 
associated  with  trees.  Its  altitudinal  range 
is  from  sea  level' at  many  localities  to  eleva- 
tions of  3500  feet  (1068  meters)  in  the 
Chaine  de  Marmelade  in  northern  Haiti 
and  3400  feet  (1037  meters)  in  the  Sierra 
de  Neiba.  Almost  all  specimens  taken  by 
myself  and  parties  were  secured  sleeping 
at  night.  \\^illiams  (1965:  2-3)  noted  that 
in  the  Monte  Cristi  region  these  lizards 
sleep  in  viny  tangles,  especially  where  there 
are  dense  "mats"  or  "curtains"  of  vines  un- 
der a  canopy.  Such  a  situation  is  ideal  in 
the  xeric  forests  in  the  Monte  Cristi  area. 
At  Las  Matas  de  Farfan,  the  lizards  were 
easily  secured  at  night  in  a  high-canopied 
cafetal,  sleeping  on  limbs,  branches,  or  on 
vines,  either  vertical  or  horizontal.  A  speci- 
men from  Morne  Calvaire  near  Petionville 
was  seen  during  the  late  morning  on  a 
mango  tree  in  an  open  pasture,  about  4  feet 
(1.2  meters)  above  the  base.  Thomas  com- 
mented in  his  field  notes  upon  a  specimen 
from  Le  Borgne  which  was  observed  8  feet 
( 2.4  meters )  above  the  ground  on  the  trunk 
of  a  tree;  this  male  led  the  pursuers  a  merry 
chase  through  a  series  of  three  trees  and 
finally  sought  refuge  in  dense  grass  on  the 
ground,  where  it  was  caught!  The  male 
from  Terrier  Rouge  was  collected  with  a 
slingshot  while  it  rested  head-down  on  the 
main  branch  of  a  large  tree  15  feet  (4.6 
meters)  above  the  ground.  South  of  Las 
Matas  de  Farfan  I  secured  a  juvenile  sleep- 
ing on  a  horizontal  vine  in  a  tree-fern 
thicket  adjacent  to  a  mountain  brook.  The 
association  of  A.  r.  ricordi  with  rivers  or 
lakes  is  certainly  fortuitous;  the  greatest 
concentrations  of  these  lizards  occur  in  such 
obviously  mesic  situations  only  because 
there  is  often  gallery  forest  restricted,  or 
limited  by  man,  to  streamsides.  However, 
such  a  situation  is  not  a  guarantee  of  secur- 
ing specimens.  In  our  effort  to  narrow  the 
previously  existing  gap  between  ricordi  and 
haleatus  in  the  northwestern  Republica 
Dominicana,  we  questioned  natives  con- 
cerning the  occurrence  of  saltacocotes  in 
the  region  along  the  gallery-forested  Rio 
Yaque     del     Norte,     which     here     passes 


through  cactus  desert.  W'e  were  assured 
that  the  lizards  indeed  occurred  there,  and 
we  were  fortimate  in  finding  a  superb  area 
of  gallery  forest  in  the  steep-sided  valley  of 
the  Rio  Guarabo,  west  of  Los  Qucmados. 
The  Guarabo  is  a  southern  affluent  of  the 
Yaque,  and  we  had  no  doubt  that  these 
splendid  hardwoods  harbored  A.  ricordi. 
But  it  was  not  until  our  fifth  nocturnal  visit 
that  a  single  subadult  was  secured,  despite 
the  attentions  of  four  collectors.  The  woods 
here  present  a  perfect  aspect  for  A.  ricordi 
— dense  and  large  trees  connected  and  in- 
terlaced with  vines  and  lianas,  all  quite 
rich  and  mesic;  still,  our  experience  indi- 
cates that,  at  least  at  the  time  of  our  visit, 
A.  ricordi  was  distinctly  uncommon  or  diffi- 
cult to  see  in  what  elsewhere  surely  would 
have  been  a  typically  simple  area  for  col- 
lection of  these  lizards.  In  this  instance, 
demands  for  at  least  one  specimen  from  this 
region  forced  persistence  which  ultimately 
yielded  the  requisite  animal.  Such  may 
well  be  the  case  in  many  otherwise  xeric 
regions,  where  A.  ricordi  is  restricted  to 
(and  perhaps  is  rare  in)  more  mesic  river- 
ine hardwood  stands. 

Specimens  examined.  HAITI:  Dcpt.  de 
rOuest,  Source  Leclerc,  Morne  Decayette 
(MCZ  65729-31);  Diquini  (MCZ  8619, 
USNM  118902,  USNM  123347,  USNM 
123988);  Port-au-Prince  (AMNH  49501); 
Petionville  (MCZ  60013-14);  Morne  Cal- 
vaire, 1  mi.  (1.6  km)  SW  Petionville,  2300 
feet  (702  meters)  (ASFS  X1711,  ASFS 
V8514,  ASFS  V9024);  Mirebalais  (MCZ 
69404);  Lancironelle,  nr.  Mirebalais  (not 
mapped)  (MCZ  68479);  Dcpt.  de  I'Arti- 
honite,  8-9  km  W  Marmelade,  3500  feet 
(1068  meters)  (ASFS  V9925);  Dcpt.  du 
Nord  Quest,  Port-de-Paix  (MCZ  63338); 
Dcpt.  du  Nord,  3  mi.  (4.8  km)  SW  Le 
Borgne  (ASFS  V10005);  2  mi.  (3.2  km) 
SW  Cap-Haitien  (ASFS  \'10766);  Ti 
Guinin,  nr.  Cap-Haitien  (not  mapped) 
(MCZ  66147-49);  8  mi.  (12.8  km)  E  Ter- 
rier Rouge  (ASFS  V10169).  RFPOBLICA 
DOMINiCANA:  Monte  Cristi  Province,  1 
km  W  Copey  (ASFS  V1269,  ASFS  V1411- 
12,  ASFS  V1470);  Laguna  de  Salodillo,  7 


106         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


km  E  Pepillo  Salcedo  (ASFS  V1413); 
Dajahon  Province,  Restauracion  (ASFS 
V18006-08);  Santiago  Rodriguez  Province, 
1.8  mi.  (2.9  km)  W  Los  Quemados,  500 
feet  (153  meters)  (ASFS  V32160);  La  Es- 
trelleta  Province,  6.7  mi.  ( 10.7  km )  E 
Hondo  Valle,  2500  feet  (763  meters) 
(ASFS  V31428);  11.0  mi.  (17.6  km)  S 
Elias  Pina,  3400  feet  (1037  meters)  (ASFS 
V31509);  San  Juan  Province,  4.9  mi.  (7.8 
km)  NW  Vallejuelo,  2400  feet  (732  meters) 
(ASFS  V31305);  6.1  mi.  (9.8  km)  S  Las 
Matas  de  Farfan,  1800  feet  (549  meters) 
(ASFS  V14562,  ASFS  V31469,  ASFS 
V31319-26);  4.1  mi.  (6.6  km)  NW  Juan  de 
Herrera,  1600  feet  (488  meters)  (ASFS 
V3139.5-99). 

Anolis  ricordi  leberi  Williams 

AnoUs    ricordii    leberi    Williams,    1965.    Breviora, 
Mus.    Comp.    Zool.,    No.    232:    4. 

Tijpe  locality.  Camp  Perrin,  Departe- 
ment  du  Slid,  Haiti;  holotype,  MCZ  80935. 

Definition.  A  subspecies  of  A.  ricordi 
characterized  by  the  combination  of  mod- 
ally  4  snout  scales  between  second  canthals, 
6  vertical  rows  of  loreal  scales,  3  scales  be- 
tween the  supraorbital  semicircles,  4/4 
scales  between  the  interparietal  and  the 
supraorbital  semicircles,  low  number  of 
vertical  dorsal  scales  (14-21;  mean  16.5), 
low  number  of  ventral  scales  ( 15-28;  mean 
20.2),  nuchal  crest  scales  usually  moderate, 
rarely  low  in  males,  usually  low,  occasion- 
ally moderate  in  females,  subocular  scales 
in  contact  with  supralabials  in  almost  50 
percent  of  the  specimens;  males  either  pale 
yellow-green  with  four  dark  saddles  and  a 
bluish  green  flank  stripe,  or  with  about 
three  longitudinal  dark  brown  lateral 
stripes,  or  simply  dark  brown,  females 
bright  green  (much  brighter  than  males), 
with  longitudinal  black  lines  indicated  and 
at  times  a  greenish  tan  middorsal  wash; 
dewlap  bright  orange  or  orange  with  an 
anterior  brown  wash  in  males,  and  dull  or- 
ange, at  times  suffused  or  marbled  with 
brown,  in  females. 

Distribution.  Known  only  from  the  vi- 
cinity of  the  type  locality  and  Marceline, 


on  the  southern  slopes  of  the  Massif  de  la 
Hotte,  between  elevations  of  1000  and  1220 
feet  (305  and  372  meters),  Dept.  du  Sud, 
Haiti. 

Discussion.  In  contrast  to  the  situation 
in  A.  r.  ricordi,  A.  r.  leberi  is  known  from 
a  long  series  of  specimens  all  from  the  same 
general  area,  at  elevations  between  1000 
feet  and  1220  feet  (305  and  372  meters). 
Williams  (1965:  6)  assigned  a  single  juve- 
nile (MCZ  38277)  from  Tardieu,  near  Pic 
Macaya,  Dept.  du  Sud,  Haiti,  to  leberi 
with  some  reservation.  This  locality  is 
northwest  of  Camp  Perrin-Marceline,  is  on 
the  northern  slopes  of  the  Massif  de  la 
Hotte,  and  is  much  closer  to  the  known 
distribution  of  the  next  subspecies  to  be 
described  below. 

The  series  of  54  A.  r.  leberi  shows  the 
following  variation.  The  largest  male 
(ASFS  X3034)  has  a  snout-vent  length  of 
147,  the  largest  female  (AMNH  98723) 
153;  both  are  from  Camp  Perrin.  Snout 
scales  at  level  of  the  second  canthal  are  ex- 
tremely variable,  and  range  between  2  and 
7;  the  mode  is  4  (23  specimens).  The  ver- 
tical loreal  rows  vary  between  5  and  8,  with 
a  mode  of  6  (26  specimens ) .  There  are  be- 
tween 1  and  4  scales  between  the  supraor- 
bital semicircles  ( mode  3 ) .  There  are  mod- 
ally  4  scales  between  the  intei-parietal  and 
the  semicircles;  4  scales  are  involved  in  64 
percent  of  the  combinations;  actual  counts 
are  3/3  (3),  3/4  (7),  4/4  (25),  4/5  (10), 
5/5  (2),  5/6  (2),  6/6  (1),  3/5  (1),  and  5/7 
( 1 ) .  Vertical  dorsals  range  between  14  and 
21  (mean  16.5),  horizontal  dorsals  between 
15  and  24  (18.0),  and  ventrals  between  15 
and  28  (20.2).  Of  39  adult  males,  30  have 
the  nuchal  crest  scales  moderate  and  nine 
have  them  low;  of  13  females,  five  have  the 
nuchal  scales  moderate  and  eight  have  them 
low.  Body  crest  scales  are  moderate  in  12 
males  and  low  in  27  males,  whereas  only 
one  female  has  the  dorsal  crest  scales  mod- 
erate and  12  have  them  low.  The  subocu- 
lars  are  separated  from  the  supralabials  by 
1  row  of  scales  in  28  specimens  and  are  in 
contact  with  the  supralabials  in  26  speci- 
mens.   A.  r.  leberi  is  the  only  population 


I 


HisPANioLAN  Giant  Angle  •  Sclnvnrfz         1U7 


that  has  sucli  a  high  incidence  (48  percent)  from  tlie  type  locahty.   The  smallest  (MCZ 

of  subocular-supralabial  contact.  83982 )  is  a  female  with  a  snout-vent  length 

Males  show  three  basic  patterns:  1)  dor-  of  52.  The  body  is  longitudinally  streaked, 
sal  ground  color  pale  yellow-green  with  but  there  are  as  yet  no  definite  longitudinal 
four  dark  brown  saddles  and  a  bluish  green  lines.  The  subadults  (  MCZ  80949-50,  a 
flank  stripe  that  is  complete;  2)  about  three  male  with  a  snout-vent  length  of  105,  and 
longitudinal  dark  brown  stripes,  the  ecu-  a  female  with  a  snout-vent  length  of  93) 
tral  one  being  the  most  prominent  and  both  show  indications  of  the  longitudinal 
complete;  3)  and  a  uniform  dark  velvety  stripes  that  are  characteristic  of  adults,  but 
brown.  In  the  two  lighter  phases,  the  eye-  the  stripes  are  better  defined  in  the  sub- 
skin  is  pale  blue,  chin  and  throats  are  dull  adult  male  than  in  the  female.  The  two 
yellow-green,  and  the  subocular  crescent  is  adult  males  and  two  adult  females  from 
pale  blue  and  very  conspicuous.  The  dew-  Marceline  agree  in  all  pattern  details  with 
laps  in  males  are  orange  (brighter  than  any  the  topotypical  series;  Marceline  and  Camp 
Macrz  and  Paul  designation)  or  orange  Perrin  are  separated  by  about  4.5  kilome- 
with  an  anterior  brown  wash.    Females  are  ters  airline. 

bright  green  dorsally  (much  brighter  than  Comparisons.     Although    both    A.    r.    ri- 

males)   with  longitudinal  black  lines  indi-  cordi  and  A.  r.  leheri  have  several  features 

cated.   There  is  a  greenish  tan  wash  on  the  in  common,  namely,  the  moderate  to  low 

upper  surface  of  the  head,  and  there  may  nuchal   and   body   crests,   the   presence   of 

be  a  greenish  tan  middorsal  zone  on  the  some  sort  of  black  body  markings,  and  a 

body.    The  dewlap  in  females  is   dull  or-  prominent   pale   subocular   crescent,    these 

ange,     often     suffused     or    marbled     with  two     subspecies     are     eminently     distinct, 

brown,  and  the  eyeskin  is  green,  paler  than  They  differ  in:    modal   number  of  second 

that  of  the  dorsum.    In  males,  the  venter  is  canthal  snout  scales   (ricordi  7,  leheri  4), 

pale  green  and  may  be  washed  with  brown  modal   number  of  loreal   rows    ( ricordi  7, 

even   in  the  green  phase,   and  in  females  leheri  6),  modal  number  of  scales  between 

the  venter  is  pale  yellow-green,  paler  than  the  interparietal   and  supraocular  semicir- 

the  bright  green  of  the  dorsum.  cles  {ricordi  5/5,  leheri  4/4),  higher  means 

In  general  aspect,  male  A.  r.  leheri  are  of  vertical  dorsal  scales  and  ventrals  (21.1, 
lineate  dorsally  and  laterally,  the  bold  dark  24.7  in  ricordi,  16.5,  20.2  in  leheri,  respec- 
longitudinal  lines  usually  interrupted  by  tively)  and  the  very  high  incidence  of  con- 
four  irregular  pale  vertical  crossbands,  tact  between  the  subocular  scales  and  the 
which  are  in  tinn  bordered  with  darker  supralabials  in  leheri  versus  the  rarity  of 
pigment.  Although  my  field  notes  indicate  this  condition  in  ricordi.  In  addition,  the 
that  there  are  about  three  longitudinal  dark  dewlap  in  male  ricordi  is  most  often  some 
stripes  in  males,  these  three  stripes  are  the  shade  of  peach  (although  the  variation  in 
result  of  modification  of  two  stripes,  of  dewlap  shades  and  colors  in  ricordi  is  read- 
which  the  more  dorsal  is  the  broader.  In  ily  acknowledged),  whereas  in  male  leheri 
many  specimens,  this  upper  flank  stripe  the  dewlap  is  orange  or  orange  with  a 
maintains  its  integrity,  but  in  many  others  brown  anterior  wash.  A  ready  hallmark  be- 
the  upper  stripe  is  hollowed  centrally,  re-  tween  the  two  subspecies  is  the  presence  of 
suiting  in  three  narrow  dark  stripes,  rather  a  pale  preauricular  blotch  in  ricordi,  a  con- 
than  two  stripes,  of  which  the  upper  is  very  dition  always  absent  in  both  sexes  of  leheri, 
broad  and  the  lower  is  narrow.  Although  with  the  result  that  instead  of  the  pale  sub- 
females  show  some  longitudinal  striping,  it  ocular  crescent's  being  incoi-poratcd  into  a 
is  much  less  conspicuous  than  in  males,  postlabial  line  or  preauricular  blotch  as  it 
Male  throats  are  immaculate,  whereas  fe-  often  is  in  ricordi,  it  is  a  bold  and  contrast- 
male  throats  are  suffused  with  dark  green.  ing  pattern  element. 

There  are  three  juveniles  and  subadults  Remarks.     All  Camp  Perrin  specimens  of 


108         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


A.  r.  leheri  were  collected  by  natives  and 
thus  I  have  no  precise  knowledge  of  the 
habitat  nor  habits  of  this  subspecies.  Camp 
Perrin  lies  in  the  lower  southern  foothills  of 
the  high  Massif  de  la  Hotte,  at  about  1000 
feet  (305  meters),  and  the  area  in  general 
is  very  mesic  and  presumably  was  once 
well  forested,  although  now  it  supports 
cafeieres  with  a  high-canopy  hardwood 
shade  cover.  Williams  (1962:  10)  cited 
field  notes  by  A.  S.  Rand  and  J.  D.  Lazell, 
Jr.,  on  A.  r.  leheri  at  Camp  Perrin  and  Mar- 
celine;  both  accounts  involve  trees  associ- 
ated with  coffee  plantings. 

Specimens  examined.  HAITI:  Dept.  du 
Slid,  Camp  Perrin  (ASFS  X3033-,35,  ASFS 
X3038-39,  ASFS  X3041-42,  ASFS  X3182, 
AMNH  93713-36,  MCZ  80935-37,  MCZ 
S0939-42,  MCZ  80944-53,  MCZ  83982); 
Marceline  (MCZ  121115);  Marceline  area, 
ca.  1000  feet  (305  meters)  (MCZ  122269, 
MCZ  121779-80). 

Anolis  ricordi  viculus  new  subspecies 

Holotype.  USNM  193974,^  an  adult 
male,  from  Castillon,  2500  feet  (763  me- 
ters), Departement  du  Sud,  Haiti,  taken  by 
native  collector  on  2.5-26  June  1971.  Orig- 
inal number  ASFS  V25059. 

Parotypes.  ASFS  V25058,  same  data  as 
holotype;  ASFS  V25060,  same  locality  and 
collector  as  holotype,  27  June  1971;  ASFS 
V24801,  ca.  2  km  (airline)  S  Castillon, 
3500-4000  feet  (1068-1220  meters),  Dept. 
du  Sud,  Haiti,  R.  Thomas,  24  June  1971; 
ASFS  V9335,  ca.  5  km  (airline)  SE  Marche 
Leon,  2200  feet  (671  meters),  Dept.  du 
Sud,  Haiti,  native  collector,  15  March  1966; 
MCZ  119035,  Castillon,  Dept.  du  Sud, 
Haiti,  T.  P.  Webster,  A.  R.  Kiester,  and  na- 
tive collectors,  31  August  1969. 

Definition.  A  subspecies  of  A.  ricordi 
characterized  by  the  combination  of  mod- 
ally  6  snout  scales  between  the  second  can- 
thals,  7  vertical  rows  of  loreal  scales,  4 
scales  between  the  supraorbital  semicircles, 
4/4  scales  between  the  interparietal  and 
the  supraorbital  semicircles,  low  number  of 
vertical  dorsal  scales  (15-19;  mean  16.7), 
moderate  number  of  ventral  scales  ( 19-24; 


mean  21.8),  nuchal  crest  scales  usually 
moderate  but  occasionally  low  in  both 
sexes,  dorsal  body  crest  scales  low  in  both 
sexes,  subocular  scales  almost  always  sep- 
arated by  one  row  of  scales  from  suprala- 
bial  scales;  males  bright  green  dorsally 
with  powdery  pale  blue-green  lateral 
stripes,  throat  pale  green  and  unmarked, 
venter  pale  green  with  pinkish  and  yellow- 
ish suffusions,  females  dark  olive-green  to 
bright  green  with  two  purple  to  powdery 
blue-gray  flank  stripes  edged  with  dark 
brown,  lower  sides  spotted  bright  green, 
yellow-green,  or  bright  green  with  four 
bright  yellow-green  crossbands  edged  with 
black,  throat  pale  green;  dewlap  deep  yel- 
low to  orange  in  males,  dull  orange  (al- 
most brown)  to  deep  yellow  with  orange 
streaking  and  bluish  edge  in  females. 

Distribution.  Known  only  from  the  vi- 
cinity of  Castillon  on  the  northern  slopes 
of  the  Massif  de  la  Hotte  at  elevations  be- 
tween 2200  and  4000  feet  (671  and  1220 
meters)  on  the  Tiburon  Peninsula  in  Haiti; 
probably  the  subspecies  occurring  at  Tar- 
dieu  near  Pic  Macaya  (see  discussion). 

Description  of  holotype.  An  adult  male 
with  a  snout-vent  length  of  143  and  a  tail 
length  of  165  (regenerated);  snout  scales 
at  level  of  second  canthals  6,  7  vertical 
rows  of  loreal  scales,  3  scales  between  su- 
praorbital semicircles,  6/5  scales  between 
interparietal  and  supraorbital  semicircles, 
vertical  dorsals  15,  horizontal  dorsals  22, 
ventrals  20,  one  row  of  scales  between  sub- 
oculars  and  supralabials,  fourth  toe  lamel- 
lae on  phalanges  II  and  III  31,  nuchal  crest 
scales  moderate,  body  crest  scales  low;  in 
life,  bright  green  above  with  a  pair  of  lat- 
eral stripes  on  each  flank  powdery  pale 
blue-green,  the  same  color  also  on  the  face; 
throat  and  neck  pale  blue-green;  venter 
pale  green  with  pinkish  and  yellowish  suf- 
fusions; dewlap  deep  yellow,  almost  or- 
ange. 

Variation.  The  series  of  three  males  and 
three  females  shows  the  following  varia- 
tion. The  largest  male  (ASFS  V25058)  has 
a  snout-vent  length  of  148,  the  largest 
female  (ASFS  V25060)  141;  both  are  from 


HisPANioLAN  Giant  Angle  •  Schwartz 


109 


Castillon.  Snout  scales  at  level  of  the  sec- 
ond canthal  range  between  5  and  9;  the 
mode  is  6  (four,  specimens).  The  vertical 
loreal  rows  are  6  or  7,  with  a  mode  of  7 
( five  specimens ) .  There  are  be^^veen  3  and 
5  scales  between  the  supraorbital  semicir- 
cles (mode  4).  There  are  modally  4  scales 
between  the  interparietal  and  the  supraor- 
bital semicircles;  4  scales  are  involved  in  58 
percent  of  the  combinations;  actual  counts 
arc  4/4  (3),  4/5  (l),5/5  (1),  and  5/6  (1). 
X'ertical  dorsals  range  between  15  and  19 
(mean  16.7),  horizontal  dorsals  between  17 
and  27  (20.0),  and  ventrals  between  19 
and  24  (21.8).  Of  three  males,  two  have 
the  nuchal  crest  scales  moderate  and  one 
has  them  low;  the  same  situation  applies 
to  the  three  females.  All  specimens  have 
the  body  crest  scales  low.  The  suboculars 
are  usually  separated  from  the  supralabials 
by  one  row  of  scales  and  are  in  contact 
with  the  supralabials  in  one  individual  ( 17 
percent ) . 

Thomas's  field  notes  on  three  males  show 
the  variation  in  dorsal  coloration  and  pat- 
tern. The  dorsum  was  bright  green  with 
the  flank  stripes  powdery  pale  blue- green, 
this  color  occurring  also  on  the  face.  The 
throat  and  neck  were  also  pale  blue-green 
and  the  venter  was  pale  greenish  with  pink 
and  yellow  suffusions.  One  male  (ASFS 
V9335)  also  had  a  white  shoulder  patch, 
but  other  pattern  details  on  this  individual 
were  lacking  since  the  specimen  was  badly 
damaged.  The  dewlap  in  the  males  was  re- 
corded as  deep  yellow  ( almost  orange )  and 
orange  (PI.  11L6).  One  female  was  green 
to  dark  olive-green  dorsally  with  two  pur- 
ple flank  stripes,  edged  with  dark  brown, 
which  were  powdery  blue-gray  anteriorly. 
The  lower  sides  were  spotted  and  suffused 
with  bright  green  or  yellow-green.  The 
venter  was  pale  green  with  a  pinkish  wash 
in  the  pectoral  region.  The  second  female 
was  marked  in  quite  a  different  fashion, 
and  the  specimen  still  maintains  the  pattern 
after  preservation.  The  dorsum  was  bright 
green  with  four  bright  yellow-green  trans- 
verse body  bands  with  black  edges;  in  this 
specimen    longitudinal    stripes    were    also 


present  but  only  in  the  nuchal  region,  and 
the  venter,  including  the  throat,  was  pale 
green.  In  both  females,  the  dewlaps  were 
recorded  as  "very  dull  orange"  and  "deep 
yellow,  almost  brown,  anteriorly,  with  or- 
angish  longitudinal  striae,  each  edged  with 
dark  gray-green,  between  striae  pale  gray- 
green  and  most  basal  striae  greenish;  edge 
of  dewlap  grayish  (faintly  blue)." 

Comparisons.  A.  r.  viculus  is  so  very 
different  from  A.  r.  ricordi  in  both  color 
and  pattern  that  detailed  comparisons  are 
hardly  necessary.  The  black  occipital,  nu- 
chal, and  anterior  body  blotches  of  male 
ricordi  are  absent  in  male  viculus,  and  the 
longitudinally  striped  pattern  in  both  sexes 
of  viculus  does  not  occur  in  ricordi.  The 
two  subspecies  differ  also  in  scale  counts, 
as  follows:  modal  number  of  snout  scales 
at  second  canthals  ( ricordi  7,  viculus  6 ) , 
scales  between  interparietal  and  supraor- 
bital semicircles  {ricordi  5/5,  viculus  4/4), 
and  much  lower  means  of  vertical  dorsal 
and  ventral  scales  (21.1,  24.7  in  ricordi, 
16.7,  21.8  in  viculus,  respectively).  The 
two  taxa  are  similar  in  number  of  loreal 
rows,  number  of  scales  between  the  semi- 
circles, and  in  relative  frequency  of  contact 
between  the  subocular  and  supralabial 
scales. 

In  every  way,  viculus  is  much  closer  to 
leheri  than  to  ricordi.  The  basic  pattern 
elements  are  comparable  in  these  two  sub- 
species, since  both  are  lineate;  however, 
the  longitudinal  flank  stripes  in  leheri  are 
dark,  whereas  in  viculus  they  are  light;  the 
single  banded  female  viculus  is  quite  dif- 
ferent in  general  aspect  from  banded  le- 
heri. As  far  as  scale  counts  are  concerned, 
the  two  subspecies  differ  in  the  following 
manner:  modal  number  of  snout  scales  at 
second  canthals  {leheri  4,  viculus  6),  num- 
ber of  vertical  loreal  rows  {leheri  6,  vicu- 
lus 7),  and  scales  between  supraorbital 
semicircles  {leheri  3,  viculus  4).  In  mean 
number  of  vertical  dorsals  and  ventrals,  the 
two  subspecies  are  very  similar,  and  both 
have  the  4/4  condition  as  the  mode  for  the 
interparietal-semicircle  relationship. 

Discussion.     Williams   (1962:  7-8)   con- 


110         BuUetm  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


sidered  the  four  specimens  then  available 
from  the  central  portion  of  the  Tiburon 
Peninsula  as  intergrades  between  ricordi 
and  leheri.  A  few  more  specimens  have  ac- 
cumulated since  that  time;  now  there  are 
one  adult  male,  one  juvenile  male,  and  six 
adult  females  from  this  central  region,  as 
follows:  HAITI:  Dept.  du  Sud,  Pemel,  nr. 
Miragoane  (not  mapped)  ( MCZ  66015- 
16),  PaiUant,  1800  feet  (549  meters) 
(ASFS  V26535-37);  Fond  des  Negres 
(ASFS  V26254,  USNM  72631,  USNM 
72633).  As  preserved,  the  adult  male 
shows  fairly  obvious  longitudinal  streaking 
of  gray  and  dull  green,  a  few  scattered 
dark  flecks  or  small  blotches  above  the 
forelimb  insertion,  and  a  prominent  suboc- 
ular  pale  crescent.  The  adult  male  is  an 
almost  ideal  representation  of  extreme  in- 
tergradation  between  viculus  and  ricordi, 
with  both  pale  longitudinal  stripes  and 
scattered  remnants  of  the  typical  ricordi 
extensive  body  blotching.  Three  recently 
collected  females  in  life  were  green  with 
longitudinal  stripes,  which  were  delimited 
by  absence  of  black  flecking  that  occurs 
elsewhere  on  the  green  ground.  In  the 
brown  phase,  these  longitudinal  stripes  had 
a  reddish  wash.  In  all  females,  the  pale 
subocular  crescent  is  obvious  and  bold,  and 
in  one  female  (MCZ  66016)  there  is  an  ad- 
ditional preauricular  pale  area  that  resem- 
bles the  condition  in  nominate  ricordi.  I 
have  no  color  data  on  the  male  dewlaps, 
but  that  of  one  female  (ASFS  V26535)  was 
dull  yellow  distally  and  pale  blue,  smudged 
with  charcoal,  basally;  the  dewlap  scales 
were  yellow-green.  The  juvenile  male 
(snout-vent  length  79)  in  life  had  a  pat- 
tern of  longitudinal  dorsolateral  stripes  and 
dorsal  crossbands,  with  a  pale  yellow  sub- 
ocular  crescent.  I  interpret  these  lizards  as 
intergradient  between  ricordi  and  viculus. 
The  central  Tiburon  localities,  however, 
are  far  removed  from  the  known  localities 
of  viculus  (110  km)  on  one  hand  and  of 
ricordi  (70  km)  on  the  other.  Williams 
(1965:  7)  regarded  the  Fond  des  Negres 
and  Pemel  specimens  as  ricordi  X  leheri 
intergrades,  and  they  could  indeed  be  so 


interpreted.  Since,  however,  leheri  occurs 
on  the  southern  slopes  of  the  Massif  de  la 
Hotte,  and  viculus  on  the  northern  slopes 
of  that  range,  and  since  all  intergradient 
specimens  are  from  the  northeastern  re- 
gions of  the  extreme  eastern  portion  of  the 
Massif  de  la  Hotte,  it  seems  much  more 
likely  that  these  central  Tiburon  specimens 
are  intergradient  between  ricordi  and  vic- 
ulus on  geographic  grounds.  They  do  not 
disagree  with  my  concepts  of  how  inter- 
grades between  these  two  subspecies  prob- 
ably should  appear.^ 


1  Since  the  above  comments  on  the  intergradi- 
ent specimens  were  written,  Williams  has  secured 
a  series  of  28  lizards  (MCZ  132302-29)  from  St. 
Croix,  1  mi.  (1.6  km)  from  Paillant,  Dept.  du 
Sud,  Haiti,  from  this  same  general  region.  There 
are  no  color  data  on  the  specimens.  The  measure- 
ments ( in  mm )  and  scale  counts  of  these  lizards, 
combined  with  those  from  the  eight  previously 
available  soecimens,  follow.  Largest  male  ( MCZ 
132325)  155,  largest  female  (ASFS  V26535)  148. 
Snout  scales  at  second  canthals  4-9  (mode  6); 
loreal  rows  4-8  ( mode  6 ) .  Modally  4/4  scales 
between  the  interparietal  and  the  supraorbital 
semicircles;  other  counts:  3/3  (2),  3/4  (2),  4/5 
(4),  5/5  (11),  5/6  (1),  3/5  (1);  4  scales  are  in- 
volved with  50  percent  of  the  combinations.  Ver- 
tical dorsals  range  between  14  and  21  (mean 
17.0),  horizontal  dorsals  between  15  and  24 
(18.3),  and  ventrals  between  16  and  31  (21.4). 
Of  the  males,  four  have  the  nuchal  crest  scales 
moderate  and  15  have  them  low;  of  the  females, 
one  has  the  nuchal  crest  scales  moderate  and  15 
have  them  low.  Body  scales  are  low  in  all  adult 
specimens.  The  suboculars  are  modally  separated 
from  the  supralabials  by  one  row  of  scales  and 
are  in  contact  with  the  supralabials  in  seven  liz- 
ards ( 19  percent). 

In  scale  counts,  the  entire  series  is  much  closer 
to  viculus  than  to  ricordi;  however,  in  some  char- 
acteristics, the  series  is  closer  to  lebcri  or  to  the 
subspecies  yet  to  be  described  from  extreme  south- 
eastern Haiti.  In  fact,  comparison  of  the  scale 
counts  shows  that  there  is  little  resemblance  be- 
tween the  modes  and  means  between  these  geo- 
graphically intermediate  specimens  and  nominate 
ricordi,  and  as  a  whole  they  seem  much  more 
closely  allied  to  one  of  the  other  Tiburon  subspe- 
cies. 

The  males  in  the  St.  Croix  series  are  variable 
in  pattern  but  none  shows  any  clear-cut  dark 
blotching,  typical  of  A.  r.  ricordi.  Some  males 
are  more  or  less  unicolor  (medium  brown  as  pre- 
served), whereas  others  have  longitudinal  stripes, 
alternating  light  and  dark,  with  usually  one  broad 


HisPANioLAx  Giant  Angle  •  Schtvartz 


111 


There  remains  one  other  specimen  from 
the  distal  portion  of  the  Tibnron  Peninsnhi; 
this  is  a  jnvenile  male  ( MCZ  38277)  with 
a  snont-vent  length  of  7(S,  from  Tardieu, 
near  Pic  Macaya,  collected  by  P.  J.  Darling- 
ton. It  is  presently  dull  brown,  but  there 
are  clear  indications  of  black-edged  dorsal 
crossbands  that  closely  resemble  the  condi- 
tion in  one  of  the  female  paratypes  of  vi- 
culus.  Tardieu  is  presently  unlocatable  on 
modern  maps,  but  Darlington  has  indicated 
to  Williams  that  this  place  lies  just  to  the 
north  of  Pic  Macaya,  and  thus  rather  close 
to  Castillon.    Since  there   are   no  juvenile 


dark  stripe  along  the  upper  sides  and  most  prom- 
inent. One  male  has  extensive  dark  brown  body 
markings,  vertically  oriented  and  alternating  with 
paler  tannish  areas  to  give  a  more-or-less  verti- 
cally barred  appearance.  The  pale  subociilar 
crescent  is  very  obvious  in  all  males,  and  there  is 
no  indication  of  a  pale  preauricular  area. 

The  females  are  undistinguished.  Most  are 
more  or  less  solid  green  with  some  scattered  paler 
green  scales  to  give  a  beadwork  effect  dorsally, 
but  there  are  also  indications  in  some  specimens 
of  longitudinal  paler  areas  to  give  a  somewhat 
longitudinally  lined  appearance.  As  in  the  males, 
the  subocular  pale  crescent  is  obvious,  and  there 
may  be  a  weakly  differentiated  pale  preauricular 
area. 

This  newly  collected  series  of  A.  ricordi  is  puz- 
zling. The  entire  lot  is  so  like,  in  general  aspect, 
specimens  of  leberi  and  viculus  (and  the  yet-to- 
be-named  subspecies  in  southwestern  Haiti )  and 
shows  so  little  tendency  toward  A.  r.  ricordi  that 
it  is  difficult  to  interpret  them  as  intergradient  be- 
tween ricordi  and  viculus.  The  adult  male  (MCZ 
66015)  noted  above  in  the  body  of  the  text  is 
from  "Pemel,  near  Miragoane,"  a  site  that  is  un- 
locatable on  modern  maps.  Pemel  may  be  "near 
Miragoane"  in  only  the  very  broadest  sense.  Spec- 
imens that  are  known  to  have  been  taken  in  the 
Miragoane-Paillant  area  show  little  evidence  of 
intergradation  between  vicidus  and  ricordi,  and 
are  much  closer  to  the  former  subspecies. 

Everything  considered,  I  strongly  suspect  that 
with  additional  collecting  on  the  central  Tiburon 
Peninsula  it  is  probable  that  two  taxa  will  be 
found  to  occur  here  in  sympatry  and  without 
wide  areas  of  intergradation,  or  that  ricordi-styled 
anoles  occur  close  to  ( but  do  not  intergrade  with ) 
leberi-styled  anoles.  The  evidence  at  the  moment 
is  far  from  unequivocal  that  ricordi  and  viculus 
intergrade  in  this  area.  Only  much  additional 
collecting  along  the  Tiburon  Peninsula  will  reveal 
the  actuality  of  the  relationships  of  A.  ricordi  with 
its  southeastern  relatixes. 


viculus,  I  have  no  concepts  of  their  appear- 
ance; on  geographic  grounds,  however,  I 
have  little  doubt  that  the  Tardieu  male  is 
representatixe  of  viculus  rather  than  of 
leberi. 

Remarks.  It  may  seem  remarkable  that 
there  should  be  two  distinctive  subspecies 
of  A.  ricordi  in  such  close  geographical 
proximity;  Castillon  and  Marceline  are  sep- 
arated by  only  29  kilometers  airline,  and  if 
the  Tardieu  specimen  is  viculus,  then  the 
distance  between  the  localities  for  the  two 
subspecies  is  even  shorter.  However,  be- 
tween Castillon  and  Marceline  lies  the  high 
ridge  of  the  La  Hotte,  including  the  cul- 
minating peak  of  that  range,  Pic  Macaya, 
with  an  elevation  of  7698  feet  (2347  me- 
ters). Such  high  and  rugged  country  is 
probably  ecologically  unsuitable  for  A.  ri- 
cordi, and  the  northern  and  southern  pop- 
ulations associated  with  the  La  Hotte  have 
differentiated  because  of  isolation  caused 
by  the  intervening  massif. 

The  known  altitudinal  range  of  A.  r.  vic- 
ulus is  between  2200  and  4000  feet  (671 
and  1220  meters).  The  Castillon  area,  ac- 
cording to  Richard  Thomas,  is  generally 
mesic  but  much  of  the  original  forest  has 
been  cut.  Still,  enough  trees  and  ravine 
woods  remain  to  offer  haven  for  such  a 
tolerant  and  adaptable  species  as  A.  ricordi. 
One  female  from  south  of  Castillon  was 
taken  by  Thomas  on  the  trunk  of  a  large 
tree  about  5  feet  (1.5  meters)  above  the 
ground;  all  other  ASFS  specimens  were  se- 
cured by  natives. 

The  name  vicidus  is  from  the  Latin  for 
"hamlet"  or  "small  village"  in  allusion  to 
Castillon,  the  type  locality. 

Anol'is  ricordi  subsolanus  new  subspecies 

Holotype.  MCZ  130270,  an  adult  male, 
from  Source  Carroye,  near  Saltrou,  Depart- 
ement  de  I'Ouest,  Haiti,  one  of  a  series  col- 
lected by  George  Whiteman  in  March  1972. 

Paratypes.  MCZ  130264-69,  MCZ 
130271-77,  same  data  as  holotype;  MCZ 
69405,  nr.  Saltrou,  Dept.  de  I'Ouest,  G. 
Whiteman,  summer  1962. 

Definition.     A   subspecies   of  A.    ricordi 


112         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


characterized  by  the  combination  of  mod- 
ally  5  snout  scales  between  second  canthals, 
5  vertical  rows  of  loreal  scales,  3  scales  be- 
tween the  supraorbital  semicircles,  4/4 
scales  between  the  interparietal  and  the 
supraorbital  semicircles,  moderate  number 
of  vertical  dorsal  scales  (16-21;  mean 
17.3),  moderate  number  of  ventral  scales 
(18-27;  mean  21.0),  nuchal  crest  scales 
rarely  moderate,  usually  low  in  males,  low 
in  females,  subocular  scales  always  sepa- 
rated from  supralabials  scales  by  one  row 
of  scales;  males  vaguely  lineate  dorsally 
with  two  broad  lateral  grayish  flank  stripes, 
or  with  three  paler  ( green  in  life? )  cross- 
bands;  females  like  males,  or  heavily 
blotched  with  black  laterally  and  on  the 
occiput,  the  black  lateral  markings  in  the 
areas  that  are  elsewhere  occupied  by 
the  gray  lateral  flank  stripes;  a  pale  suboc- 
ular crescent  present  and  prominent  but  no 
pale  preauricular  blotch;  dewlap  color  un- 
known. 

Distribution.  Known  only  from  the  re- 
gion about  Saltrou,  in  extreme  southeastern 
Haiti,  but  see  discussion  below. 

Description  of  holotype.  An  adult  male 
with  a  snout-vent  length  of  144  mm  and  a 
tail  length  of  209  mm  ( regenerated ) ;  snout 
scales  at  level  of  second  canthals  4,  6  ver- 
tical rows  of  loreal  scales,  3  scales  between 
supraorbital  semicircles,  3/4  scales  between 
the  interparietal  and  the  supraorbital  semi- 
circles, vertical  dorsals  18,  horizontal  dor- 
sals 17,  ventrals  19,  one  row  of  scales  be- 
tween suboculars  and  supralabials,  fourth 
toe  lamellae  on  phalanges  II  and  III  33, 
nuchal  crest  scales  low,  body  crest  scales 
low;  as  preserved,  dorsum  dull  dark  brcwn 
with  three  prominent  blue-green  cross- 
bands,  more  or  less  confluent  middorsally, 
and  outlined  in  dark  brown  to  black;  top 
of  head  brown,  paler  than  sides;  throat 
greenish,  dewlap  dull  gray;  belly  dark 
gray,  underside  of  hindlimbs  green;  tail 
brown. 

Variation.  The  holotype  and  paratypic 
series  are  composed  of  10  males  and  five 
females.    The  largest  male  (MCZ  130274) 


has  a  snout-vent  length  of  152,  the 
largest  female  (MCZ  69405)  150;  the  male 
is  a  topotype,  the  female  is  from  near  Sal- 
trou. Snout  scales  at  level  of  the  second 
canthals  range  between  4  and  7;  the  mode 
is  5  (six  specimens).  The  vertical  loreal 
rows  vary  between  5  and  7,  with  a  mode  of 
5  (eight  specimens).  There  are  between  2 
and  4  scales  between  the  supraorbital  semi- 
circles (mode  3).  There  are  modally  4 
scales  between  the  intei-parietal  and  the 
supraorbital  semicircles;  4  scales  are  in- 
volved in  58  percent  of  the  combination; 
actual  counts  are  3/3  ( 1),  3/4  (3),  4/4  (4), 
4/5  (3),  5/5  (2),  and  5/6  (2).  Vertical 
dorsals  range  between  16  and  21  (mean 
17.3),  horizontal  dorsals  between  14  and  23 
(17.1),  and  ventrals  between  18  and  27 
(21.0).  Of  10  males,  two  have  the  nuchal 
crest  scales  moderate  and  eight  have  these 
scales  low;  all  five  females  have  the  nuchal 
crest  scales  low.  All  specimens  have  the 
body  crest  scales  low.  In  all  specimens  the 
subocular  scales  are  separated  from  the  su- 
pralabials by  one  row  of  scales. 

I  have  no  color  notes  in  life  nor  have  I 
seen  live  specimens  of  A.  r.  suhsolanus. 
Consequently,  my  comments  on  pattern  in 
this  subspecies  are  based  solely  upon  pre- 
served material.  In  the  series  of  males  and 
females,  each  sex  shows  two  basic  patterns. 
The  more  common  is  a  pair  of  longitudinal 
flank  stripes,  the  upper  being  broader,  usu- 
ally dull  grayish  in  contrast  to  a  greenish 
ground  color.  In  two  specimens  (one  male 
and  one  female;  MCZ  130267  and  MCZ 
69405)  these  stripes  are  very  prominent 
and  black;  although  they  no  longer  have 
their  integrity  in  the  female,  they  are  still 
very  obvious.  In  addition,  in  the  female 
there  is  black  pigment  in  the  occipital  re- 
gion. A  pale  subocular  crescent  is  present 
in  all  specimens  and  is  usually  very  con- 
spicuous. In  two  specimens  (the  holotypic 
male  and  a  female— MCZ  130266)  the  dor- 
sal pattern  consists  of  three  transverse 
crossbands  that  are  green,  more  or  less 
fused  middorsally,  and  outlined  with  black 
or  dark  brown.  Many  females  show  the  lat- 


HisPANioLAN  Giant  Angle  •  Schtvartz         113 


eral  flank  stripes  much  less  clearly  than  do 
the  males,  but  usually  the  stripes  are  at 
least  indicated.    • 

Comparisons.  In  general  aspect,  sub- 
sola  nus  much  more  closely  resembles  far 
western  leheri  and  vicuhis  than  geographi- 
cally closer  ricordi.  The  latter  subspecies, 
however,  occurs  on  the  nortliern  side  of  the 
Massif  de  la  Selle,  whereas  the  localities  for 
suhsolamis  are  to  the  south  of  that  range. 
Since  I  do  not  know  the  coloration  in  life 
of  suhsolamis,  I  am  unable  to  compare  its 
pigmentation  with  that  of  the  other  subspe- 
cies. The  presence  of  both  longitudinally 
striped  and  transversely  barred  specimens 
in  suhsolamis  suggests  its  affinity  with  leheri 
and  vicuhis.  A.  r.  suhsolamis  differs  from 
A.  r.  ricordi  in  that  the  latter  has  (in  its 
southern  populations)  dark  anterior  mark- 
ings on  the  occiput  and  above  the  forelimb 
insertions,  whereas  these  markings  are  ab- 
sent in  subsolanus.  Additionally,  southern 
ricordi  are  patternless  green,  whereas  sub- 
solaiius  females  are  longitudinally  lined  and 
may  have  heavy  dark  anterior  markings 
(somewhat  like  male  A.  r.  ricordi).  At  the 
time  of  Williams's  review  of  A.  ricordi 
(1965:  2),  there  was  but  a  single  A.  ricordi 
from  the  Saltrou  region;  by  chance,  this 
specimen  (MCZ  69405)  is  the  heavily 
marked  female  upon  which  I  commented 
above.  Although  Williams  (loc.  cit.)  con- 
sidered it  a  male,  it  lacks  enlarged  postanal 
scales  and  a  tail  "fin,"  and  it  is  a  female. 
Since  female  A.  r.  ricordi  lack  dark  anterior 
markings,  this  female  is  really  quite  differ- 
ent from  females  of  the  northern  subspe- 
cies. 

From  the  western  subspecies  leheri  and 
viculus,  suhsolamis  difi^ers  meristically  in 
the  following  ways.  From  leheri,  suhso- 
lanus  differs  in  having  5  versus  4  snout 
scales  at  the  second  canthal,  5  versus  6  ver- 
tical loreal  rows,  higher  means  in  vertical 
dorsal  scales  and  ventral  scales,  and  also 
lacks  specimens  that  have  the  suboculars  in 
contact  with  the  supralabials  ( leheri  has  48 
percent  of  the  specimens  with  this  condi- 
tion).   From  viculus,  suhsolanus  differs  in 


having  5  versus  6  snout  scales  at  second 
canthal,  5  versus  7  vertical  loreal  rows,  3 
versus  4  scales  between  the  supraorbital 
semicircles,  and  higher  means  of  vertical 
dorsal  scales  and  ventral  scales.  In  addition 
to  the  pattern  differences  noted  above 
which  differentiate  suhsolanus  from  nomi- 
nate ricordi,  suhsolanus  has  5  versus  7  snout 
scales  at  the  second  canthals,  5  versus  7 
vertical  loreal  rows,  3  versus  4  scales  be- 
tween the  supraorbital  semicircles,  4/4  ver- 
sus 5/5  scales  between  the  interparietal  and 
the  supraorbital  semicircles,  and  lower 
means  in  vertical  dorsal  scales  and  ventral 
scales. 

Remarks.  I  am  once  more  hampered  in 
my  interpretation  of  suhsolanus  by  the 
large  distributional  gap  between  its  two 
stations  and  any  other  stations  for  A.  ricordi 
to  the  west.  The  absence  of  specimens 
from  the  southern  coast,  from  such  well- 
known  areas  as  Jacmel  and  Aquin,  is  truly 
puzzling.  The  nearest  locality  to  suhsolanus 
along  the  Tiburon  Peninsula  is  Fond  des 
Negres  [ricordi  X  viculus),  some  120  kilo- 
meters to  the  west.  Still  further,  the  area 
known  to  be  occupied  by  A.  r.  leheri  lies 
some  205  kilometers  to  the  west,  near  the 
peninsula's  tip.  Known  stations  for  A.  r.  ri- 
cordi are  very  much  closer  (40  kilometers) 
but  lie  to  the  north  of  the  Massif  de  la 
Selle.  Closer  even  than'  any  of  these  is 
harahonae;  harahonae  and  suhsolanus  are 
known  in  this  region  for  localities  separated 
by  about  11  kilometers  (see  comments  be- 
low), but  there  is  no  evidence  of  intergra- 
dation  between  these  two  taxa. 

It  is  perhaps  pertinent  that  leheri,  vicu- 
lus, and  suhsolanus  all  seem  closer  in  most 
characteristics  to  each  other  than  they  do 
to  nominate  ricordi.  If  it  were  not  for  the 
specimens  that  I  interpret  as  intergradient 
between  ricordi  and  viculus  in  the  Mira- 
goane-Paillant-Fond  des  Negres  region,  I 
would  be  very  tempted  to  consider  these 
three  taxa  as  a  species  distinct  from  A.  ri- 
cordi. Much  additional  material  from  along 
the  Tiburon  Peninsula  will  perhaps  show 
that  my  interpretation  is  wrong. 


114         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


The  name  subsolamis  is  from  the  Latin 
for  "eastern,"  in  alhision  to  the  occurrence 
of  this  subspecies  in  southeastern  Haiti. 
The  precise  areas  where  suhsolanus  occurs 
are  a  matter  of  question.  I  am  unable  to 
locate  Source  Carroye  on  any  modern  map. 
Williams  advised  me  that  Source  Carroye 
is  very  near  Thiotte  (according  to  the  col- 
lector, "Source  Carroye  is  located  northeast 
direction  and  about  V2  mile  from  the  main 
road  after  you  leave  the  place  of  the 
'marche',"  that  market  being  at  Thiotte). 
The  elevation  of  Thiotte  is  about  900  me- 
ters. The  lone  specimen  from  "near  Sal- 
trou"  also  poses  the  problem  of  just  how 
"near"  this  specimen  was  taken  to  Saltrou 
itself.  Any  information  on  details  of  local- 
ities or  elevations  of  this  and  other  speci- 
mens taken  along  the  Dominico-Haitian 
border  are  mandatory.  The  distance  be- 
tween the  Thiotte  locality  for  suhsolanus 
and  the  Pedernales  specimens  of  harahonae 
is  about  11  kilometers.  It  is  especially  per- 
tinent that  harahonae  is  not  known,  along 
the  Dominico-Haitian  border,  from  the  low- 
lands (where,  incidentally,  Anolis  coeles- 
tinus  is  called  saltacocote  by  the  natives), 
but  that  harahonae  occurs  here  as  an  in- 
habitant of  mesic  riverine  woods  at  an  ele- 
vation of  600  feet  ( 183  meters ) . 

Anolis  barahonoe  Williams 

Anolis  ricordii  harahonae  Williams,    1962.     Brevi- 
ora,  Mus.  Comp.  Zool.,  No.  155:  8. 

Ttjpe  locality.  Polo,  Valle  de  Polo,  Bara- 
hona  Province,  Repiiblica  Dominicana; 
holotype,  MCZ  43819. 

Defiyiition.  A  giant  species  of  Hispanio- 
lan  Anolis  characterized  by  the  combina- 
tion of  moderate  size  (males  to  158  mm, 
females  to  148  mm  snout-vent  length), 
snout  scales  at  level  of  second  canthal 
scales  2  to  5  (mode  4),  vertical  loreal  rows 
2  to  5  (mode  6),  scales  between  supraor- 
bital semicircles  1  to  4  (mode  2),  inteipa- 
rietal  scale  separated  from  supraorbital 
semicircles  modally  by  4  scales,  vertical 
dorsal  scales  generally  small  ( 15  to  34  in 
standard-distance),  ventral  scales  relatively 
small  (17  to  29  in  standard-distance),  nu- 


chal crest  scales  in  both  sexes  rarely  high, 
usually  moderate  to  low,  dorsal  body  crest 
scales  rarely  moderate,  usually  low,  suboc- 
ular  scales  rarely  in  contact  with  suprala- 
bial  scales;  dorsal  body  coloration  basically 
lichenate  gray-green,  grays,  to  browns  and 
black,  giving  a  blotched  effect  that  also  oc- 
curs in  even  the  smallest  juveniles,  and 
rarely  (only  in  juveniles)  with  any  indica- 
tion of  transverse  crossbars,  or  solid  brown 
to  grayish  with  faintly  bluish  white  dark- 
edged  ocelli;  dewlap  pale  yellow  to  peach 
in  males,  pale  yellow  to  pale  peach  in  fe- 
males; pale  subocular  crescent  absent  in 
adults  but  indicated  in  juveniles  by  a  pale 
subocular  spot. 

Distrihution.  The  Sierra  de  Baoruco 
and  associated  lowlands  on  the  Peninsula 
de  Barahona,  Repiiblica  Dominicana,  in- 
cluding (probably)  the  semi-xeric  forests 
of  the  lowlands  south  of  the  Sierra  de 
Baoruco  and  southern  Haiti;  altitudinal  dis- 
tribution from  sea  level  to  2600  feet  (793 
meters)  northeast  of  Las  Auyamas,  Bara- 
hona Province. 

Anolis  harahonae  harahonae  Williams 
Type  locality.     Polo,  Valle  de  Polo,  Bara- 
hona Province,  Repiiblica  Dominicana. 

Definition.  A  subspecies  of  A.  hara- 
honae characterized  by  the  combination  of 
modally  4  snout  scales  between  second  can- 
thai  scales,  4  vertical  rows  of  loreal  scales, 
2  scales  between  the  supraorbital  semicir- 
cles, 4/4  scales  between  the  interparietal 
and  the  supraorbital  semicircles,  relatively 
low  number  of  vertical  dorsal  scales  ( 15- 
23;  mean  17.2),  high  number  of  ventral 
scales  (17-29;  mean  22.1),  nuchal  crest 
scales  moderate  to  low,  body  crest  scales 
rarely  moderate,  usually  low,  subocular 
scales  usually  separated  from  supralabial 
scales  by  one  row  of  scales,  both  sexes  and 
juveniles  patterned  with  varying  shades  of 
gray-green,  grays,  browns  and  black,  giv- 
ing a  lichenate  blotched  effect;  juveniles 
with  vague  indications  of  three  transverse 
gray  bands  but  that  pattern  only  very 
rarely  even  indicated  in  adults;  dewlap 
pale  yellow  to  pale  peach  in  both  sexes,  the 


HisPANiOLAN  Giant  Angle  •  Schtvartz         115 


female  dewlap  suffused  with  gray  basally; 
pale  subocular  crescent  absent  in  adults  but 
indicated  by  a  clear  white  subocular  spot  in 
juveniles  and  subadults. 

Discussion.  A.  b.  harahonae  has  a  rela- 
tively circumscribed  range  in  the  Sierra  de 
Baoruco  in  the  southeastern  Republica  Do- 
minicana.  Until  our  1971  collections,  the 
taxon  had  been  known  only  from  the  east- 
ern portion  of  that  massif,  but  two  speci- 
mens taken  13.0  mi.  ( 20.8  km )  N  of  Peder- 
nales  along  the  Dominico-Haitian  border 
are  unquestionably  A.  harahonae.  These 
individuals  differ  slightly  from  more  east- 
ern specimens  of  A.  h.  harahonae  in  colora- 
tion, but  they  are  so  close  to  the  nominate 
subspecies  that  for  the  moment  I  have  no 
hesitancy  in  regarding  them  as  that  taxon. 

The  series  of  33  specimens  of  A.  h.  hara- 
honae shows  the  following  variation.  The 
largest  males  (ASFS  V29722,  MCZ  125504) 
have  snout-vent  lengths  of  158,  the  largest 
female  (AMNH  50256)  148;  the  males  are 
from  north  of  Pedernales  and  near  Polo, 
and  the  female  is  from  Barahona.  Snout 
scales  at  level  of  second  canthals  vary  be- 
tween 2  and  5;  the  mode  is  4  (18  speci- 
mens). The  vertical  loreal  rows  vary  be- 
tween 5  and  8,  with  a  mode  of  6  (11 
specimens).  There  are  between  1  and  4 
scales  between  the  supraorbital  semicir- 
cles (mode  2).  There  are  modally  4  scales 
between  the  inteiparietal  and  the  supraor- 
bital semicircles;  4  scales  are  involved  with 
58  percent  of  the  combinations;  actual 
counts  are  3/3  (3),  3/4  (6),  4/4  ( 13),  4/5 
(3),  and  5/5  (5).  Vertical  dorsals  range 
between  15  and  23  (mean  17.2),  horizontal 
dorsals  between  15  and  24  (18.2),  and  ven- 
trals  between  17  and  19  (22.1).  Of  16 
males,  seven  have  the  nuchal  crest  scales 
moderate  and  nine  have  these  scales  low; 
of  10  females,  three  have  these  scales  mod- 
erate and  seven  have  them  low.  Body  crest 
scales  are  moderate  in  one  male  and  low  in 
15,  whereas  all  10  females  have  the  body 
crest  scales  low.  The  subocular  scales  are 
separated  from  the  supralabial  scales  in  32 
of  33  specimens  (3  percent). 

Exclusive  of  the  male  and  female  from 


north  of  Pedernales,  eastern  specimens  of 
A.  /;.  harahonae  are  lichenate  or  blotclied 
with  gray-green,  grays,  browns,  and  black 
in  a  random  pattern,  although  occasional 
individuals  show  remnants  of  the  slightly 
more  obviously  banded  condition  of  the 
juveniles.  No  specimen  has  been  recorded 
in  the  field  as  being  bright  green,  and  in 
general  the  tones  of  green  in  harahonae 
are  dull  and  grayish.  Some  specimens  (es- 
pecially ASFS  V30921,  a  male)  were  re- 
corded as  being  gray,  heavily  blotched  with 
black,  and  thus  without  any  green  tints 
whatsoever.  The  dewlap  color  in  males 
varies  between  pale  peach  and  peach,  and 
in  females  between  pale  peach  and  yellow. 

The  Pedernales  specimens  were  recorded 
in  life  as  being  dark  brown  to  gray  dorsally, 
obscurely  banded  with  tannish.  The  heads 
were  tan  above,  the  eyeskin  pale  gray,  and 
the  female  had  the  upper  surfaces  of  all 
limbs  banded  green  and  dark  brown.  The 
most  noteworthy  difference  between  these 
western  specimens  and  those  from  the  east- 
ern uplands  of  the  Sierra  de  Baoruco  and 
its  associated  lowlands  is  that  the  dewlaps 
in  both  sexes  were  pale  yellow,  that  of  the 
female  suffused  with  gray  basally. 

Available  juveniles  and  subadults  vary  in 
length  between  62  and  95.  The  juveniles 
are  colored  and  patterned  essentially  like 
the  adults,  except  that  three  pale  grayish 
crossbands  are  vaguely  indicated  in  most 
specimens.  These  bands  are  quite  indis- 
tinct and  much  obscured  by  the  lichenate 
harahonae  pattern.  Some  juveniles  were  re- 
corded as  being  banded  and  mottled  pale 
gray,  dull  pea-green,  and  black,  witli  a 
black  nuchal  patch  and  a  white  nuchal 
crescent  on  each  side,  whereas  others  were 
recorded  as  crossbanded  gray  and  dusky, 
with  some  greenish  on  the  lips,  and  the  tails 
banded  gray  and  dusky  to  cream. 

Remarks.  Specimens  secured  by  myself 
and  parties  have  all  been  taken  in  wooded 
situations,  between  elevations  of  600  feet 
and  2600  feet  ( 183  and  793  meters).  River- 
ine woods  and  the  large  shade  trees  in  the 
upland  cafetales  of  the  Sierra  de  Baoruco 
offer  optimum  habitat  for  the  species.   Both 


116         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


adults  and  juveniles  were  secured  sleeping 
at  night;  in  general,  the  juveniles  sleep 
lower  on  shrubs  and  low  trees,  whereas 
adults  sleep  higher  (up  to  15  feet — 4.6  me- 
ters) on  limbs,  branches,  and  woody  vines. 
At  night,  despite  the  absence  of  bright 
greens  in  the  coloration,  the  lizards  are 
quite  obvious  because  their  pale  grayish 
hues  contrast  to  the  adjacent  greenery.  All 
ages  of  A.  h.  harahonae  sleep  exposed,  as 
do  other  Hispaniolan  giant  anoles.  The 
pair  from  13.0  mi.  N  Pedernales  were  se- 
cured in  rich  riverine  woods  at  an  eleva- 
tion of  600  feet  ( 183  meters ) ;  this  is  purely 
a  gallery  forest  situation,  since  in  this  re- 
gion the  open  slopes  are  clad  in  Acacia  for- 
est or  dry  scrubby  woodlands,  whereas 
rivers  and  creeks  support  much  more  luxu- 
riant arboreal  growth. 

Almost  all  localities  for  A.  h.  harahonae 
are  in  the  highlands.  However,  the  lizard 
presumably  occurs  in  coastal  forested  re- 
gions as  well.  There  are  specimens  from 
the  city  of  Barahona  ( which  is  coastal )  and 
from  halfway  between  Enriquillo  and 
Oviedo,  which  is  presumed  to  be  coastal  or 
nearly  so.  A  third  specimen  from  Enri- 
quillo likewise  is  presumably  from  a  coastal 
locality.  However,  in  each  of  these  cases, 
it  is  possible  that  the  lizards  were  secured 
in  the  adjacent  Sierra  de  Baoruco;  this 
mountain  range  comes  abruptly  to  the  coast 
between  Barahona  and  Enriquillo,  and  it 
would  be  a  simple  matter  to  label  speci- 
mens from  non-coastal  localities  as  having 
come  from  coastal  populated  areas.  Al- 
though negative  evidence  at  best,  we  have 
never  oiuselves  secured  A.  harahonae  along 
this  coastal  region,  and  residents  of  Bara- 
hona responded  negatively  when  ap- 
proached to  collect  this  lizard  for  us. 

A.  /;.  harahonae  is  known  from  a  locality 
(13.0  mi.  N  Pedernales)  that  is  only  (pre- 
sumably) 11  kilometers  from  a  locality 
(Thiotte)  where  A.  r.  suhsolanus  occurs. 
There  are  no  other  localities  where  these 
two  species  approach  each  other,  although, 
since  the  northern  slopes  of  the  Sierra  de 
Baoruco  are  confluent  with  the  northern 
slopes  of  the  Massif  de  la  Selle  and  its  affil- 


iates, it  is  not  unlikely  that  somewhere 
along  these  northern  reaches  A.  harahonae 
comes  into  contact  with  A.  r.  ricordi.  There 
is  no  obvious  reason  for  A.  h.  harahonae  to 
be  promptly  replaced  by  A.  r.  suhsolanus 
at  the  Dominico-Haitian  border;  the  polit- 
ical boundary  on  these  southern  slopes  is 
the  Rio  Pedernales,  a  small  stream  that 
surely  offers  no  obstacle  for  these  arboreal 
lizards.  It  follows  that  A.  h.  harahonae 
must  occur  in  southeastern  Haiti.  Thus,  as 
previously  noted,  the  accuracy  of  the  suh- 
solanus localities  is  more  than  academic.  It 
is  possible  that  in  southeastern  Haiti,  A. 
harahonae  is  a  more  lowland  lizard  and  A. 
ricordi  (suhsolanus)  occurs  on  the  higher 
and  better  forested  slopes  of  the  Massif  de 
la  Selle — the  division  may  thus  be  altitu- 
dinal  as  well  as  ecological.  The  precise  re- 
lationships between  these  two  species  re- 
main to  be  determined;  only  further 
detailed  collecting  in  extreme  southeastern 
Haiti  will  reveal  the  siutation  there.  As  far 
as  distinguishing  A.  r.  suhsolanus  from  A. 
h.  harahonae,  there  is  no  problem,  since  the 
styles  of  pattern  (and  presumably  colora- 
tion) are  so  very  different  as  to  preclude 
confusion.  If  intergradation  between  suh- 
solanus and  harahomie  occurs  (and  since  I 
here  regard  harahonae  as  a  species  distinct 
from  ricordi,  I  am  obviously  convinced  that 
it  does  not),  then  it  must  take  place  very 
quickly,  in  a  distance  of  some  11  kilome- 
ters, since  suhsolanus  and  the  Pedernales 
harahonae  are  completely  different  and 
typical  of  their  own  populations,  without 
any  indication  of  intergradation  between 
them. 

Specimens  examined.  REPUBLIC  A  DO- 
MINICAN A:  Barahona  Province,  Barahona 
(AMNH  50255-56);  14  km  SW  Barahona, 
1200  feet  (366  meters)  (ASPS  V23460-63, 
ASFS  V30263-70);  Valle  de  Polo  (MCZ 
56141,  AMNH  51235-37,  AMNH  51240, 
AMNH  51036);  nr.  Polo  (MCZ  125504-06); 
Las  Auyamas  (ASFS  V30921);  8  km  NE 
Las  Auyamas,  2600  feet  (793  meters) 
(ASFS  X9676);  Hermann's  finca,  nr.  Par- 
aiso  (AMNH  51231-33);  Enriquillo 
(AMNH  51241);  Pedernales  Province,  half- 


HisPANiOLAN  Giant  Angle  •  Schwartz 


117 


way  between  Enriquillo  and  Oviedo 
(AMNH  51230);  13.0  mi.  (20.8  km)  N 
Pedernales,  600  feet  ( 1S3  meters )  ( ASFS 
V29722-23);  locality  unkno\\ai  (AMNH 
51229). 

Anolis  borahonoe  olbocellotus 

new  subspecies 

Holotype.  MCZ  125611,  an  adult  male, 
from  13.1  mi.  (21.0  km)  SW  Enriquillo, 
Pedernales  Province,  Republiea  Domini- 
cana,  taken  by  Richard  Thomas  on  10  De- 
cember 1964.  Original  number  ASFS 
V4422. 

Definition.  A  sub.species  of  A.  hara- 
honae  characterized  by  the  combination  of 
4  snout  scales  between  second  canthal 
scales,  7  vertical  rows  of  loreal  scales,  3 
scales  between  the  supraorbital  semicir- 
cles, 4/4  scales  between  the  interparietal 
and  the  supraorbital  semicircles,  apparently 
relatively  high  number  of  vertical  dorsal 
scales  (19),  high  number  of  ventral  scales 
(26),  nuchal  crest  scales  high,  body  crest 
scales  low,  subocular  scales  separated  from 
supralabial  scales  by  one  row  of  scales, 
male  (females  unknown)  dorsal  ground 
color  nonlichenate  brown  to  grayish  with 
white  (faintly  bluish)  randomly  placed 
dark-edged  ocelli,  head  light  brown  above, 
dewlap  pale  yellow  with  a  pink  margin, 
and  a  pale  subocular  spot. 

DistriJnition.  Known  only  from  the  type 
locality,  but  presumably  distributed 
through  the  semi-arid  forests  of  the  Penin- 
sula de  Barahona  south  of  the  Sierra  de 
Baoruco  (see  discussion). 

Description  of  holotype.  An  adult  male 
with  a  snout-vent  length  of  150  and  a  tail 
length  of  265;  snout  scales  at  level  of  sec- 
ond canthal  4,  7  vertical  rows  of  loreal 
scales,  3  scales  between  interparietal  and 
supraorbital  semicircles,  vertical  dorsals  19, 
horizontal  dorsals  23,  ventrals  23,  one  row 
of  scales  between  suboculars  and  suprala- 
bials,  fourth  toe  lamellae  on  phalanges  II 
and  III  34,  nuchal  crest  scales  high,  body 
crest  scales  low;  in  life,  dorsum  brown  to 
grayish,  not  lichenate,  with  randomly  scat- 
tered   white    (faintly    bluish)    dark-edged 


ocelli  involving  from  1  to  4  scales;  venter 
white  with  gray  mottling  or  stippling;  dew- 
lap pale  yellow  with  pink  along  its  outer 
margin;  upper  surface  of  head  light  brown, 
with  large  pale  subcircular  areas  anterior 
to  the  ear  opening,  and  a  conspicuous  pale 
blotch  ])el()w  the  eye;  soles  of  hands  and 
feet  conspicuously  pale  yellow. 

Comparisons.  No  mensural  nor  meris- 
tic  characters  separate  aU)ocellatns  from 
barahoiuie.  On  the  other  hand,  the  distinc- 
tive coloration,  pattern,  and  dc^wlap  color 
of  aJhoceUatus  are  very  different  from  those 
of  harahonae,  and  the  presence  of  high  nu- 
chal crest  scales  likewise  differentiates  al- 
hocellatus  from  the  moderate  to  low  scales 
in  Imrahonae.  More  detailed  comparisons 
are  impossible,  but  certainly  aIJ)OceIl(itus  is 
quite  distinctive  when  compared  with  ])ara- 
honue. 

Discussion.  It  may  seem  foolhardy  to 
name  a  subspecies  of  A.  harahonae  from  a 
single  specimen  whose  locality  is  only  11 
kilometers  from  a  presumed  locality  for  A. 
h.  harahonae  (half  way  between  Enriquillo 
and  Oviedo).  The  holotype  of  A.  h.  alho- 
cellatus  is  that  lizard  about  which  Williams 
(1965:  4)  commented,  saying  it  "is  typical 
in  squamation  but  peculiar  in  having  very 

distinct  small  Ufi,ht  spots  on  the  flanks 

It  will  be  recalled  that  it  was  a  specimen 
from  Enriquillo  (AMNH  51241)  that 
caused  some  hesitation  when  harahonae 
was  first  described.  In  AMNH  51241  the 
pattern  was  thought  to  be  obscure  banding; 
the  present  specimen  clearly  shows  spots 
tending  to  be  vertically  aligned — a  condi- 
tion which  is  easily  transformed  into  verti- 
cal banding.  It  is  possible  that  the  ricordii 
populations  in  the  vicinity  of  Enriquillo 
consistently  show  a  distinctive  pattern 
though  charact(>ristically  harahoiuie  in 
squamation." 

The  specimen  (AMNH  51230)  from  half- 
way between  Enriquillo  and  Oviedo  is  a 
young  male  with  a  snout-vent  length  of 
121.  Since  this  lizard  presumably  came 
from  the  lowlands  of  the  Peninsula  de 
Barahona,  it  might  logically  be  expected  to 
be  alhocellatiis.   However,  the  lizard  is  now 


118         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


drab  patternless  brown,  and  there  are  no 
indications  that  it  was  ever  spotted.  Pre- 
sumably albocellatus  and  harahonae  inter- 
grade  between  Enriqiiillo  (which  Hes  at 
the  extreme  southeastern  corner  of  the  Si- 
erra de  Baoruco)  and  Oviedo  (which  Hes 
well  down  on  the  Peninsula  de  Barahona). 
Several  facts  have  prompted  my  naming 
this  lonely  specimen.  First,  I  have  exam- 
ined the  Enriquillo  specimen  noted  by  Wil- 
liams, and,  although  it  shows  some  indica- 
tion of  vertical  crossbars,  they  are  not  any 
more  conspicuous  than  those  in  some  more 
recently  taken  A.  /;.  harahoruie  from  the 
Baoruco  highlands  (Williams  examined 
only  17  harahoime  at  the  time  of  its  original 
description;  I  have  studied  almost  twice 
this  number).  Secondly,  the  xeric  to  semi- 
arid  region  south  of  the  Sierra  de  Baoruco 
has  come  to  be  known  as  an  area  of  local 
differentiation  at  the  subspecific  level  for  a 
variety  of  reptiles;  this  alone  is  no  reason 
for  naming  albocellatus,  of  course.  Thirdly, 
although  since  1964  when  the  holotype  was 
collected  both  I  and  others  have  spent  con- 
siderable time  on  the  Peninsula  de  Bara- 
hona and  in  the  vicinity  of  Oviedo,  we  have 
never  seen  or  secured  another  A.  harahonae 
in  this  region.  In  September  1966,  the  very 
severe  hurricane  Inez  passed  directly  across 
the  Peninsula.  What  had  once  been  high- 
canopied  semi-arid  forest  (as  at  Oviedo) 
has  been  either  totally  destroyed  or  been 
reduced  (by  1969)  to  a  landscape  of  bare 
snags  with  some  leafy  growth  just  now  be- 
ginning to  appear  but  at  a  much  lower 
canopy-level  than  previously.  The  changes 
between  the  Oviedo  area  in  1964  and  1969 
are  so  massive  that,  upon  my  first  visit  there 
after  Inez,  I  was  unable  to  orient  myself  in 
reference  to  our  older  collecting  localities! 
Certainly  this  entire  region  has  suffered 
greatly,  and,  with  the  destruction  of  trees, 
it  seems  reasonable  to  assume  that  A.  hara- 
honae has  suffered  equally.  The  population 
may  never  have  been  high,  since  such  semi- 
arid  woods  are  not  at  all  optimal  habitat 
for  any  of  the  Hispaniolan  giant  anoles,  and 
the  destruction  of  the  habitat  must  surely 


have  affected  A.  h.  albocellatus  adversely. 
Since  persistent  visits  to  this  area  have 
yielded  no  new  material,  and  since  the  liz- 
ard may  presently  be  very  rare,  I  have  de- 
cided upon  the  present  course  rather  than 
wait  in  hope  for  someone  to  secure  a  sec- 
ond ( or  more )  lizard. 

Remarks.  The  Peninsula  de  Barahona 
has  been  shown  to  have  distinctive  subspe- 
cies (or  even  species)  of  a  variety  of  rep- 
tiles. Species  that  have  described  endemic 
subspecies  south  of  the  Sierra  de  Baoruco 
include:  Sphaerodacttjhis  difficilis  Barbour, 
Leiocephalus  Imrahonensis  Schmidt,  Am- 
eiva  chrysolaema  Cope,  Ameiva  lineolata 
Dumeril  and  Bibron,  Arnphisbaena  gona- 
vensis  Cans  and  Alexander,  and  Dromicus 
parvifrons  Jan.  Endemic  Peninsula  de 
Barahona  species  are:  Anolis  longitibialis 
Noble,  Typhlops  sijntherus  Thomas,  Lep- 
totyphlops  pyrites  Thomas,  and  Uromacer 
ivetmorei  Cochran.  Only  one  amphibian, 
Eleutherodactylus  alcoae  Schwartz,  is  re- 
stricted to  the  Peninsula.  To  the  former 
list  can  now  be  added  Anolis  harahonae. 
The  eastern  half  of  the  Peninsula,  although 
xeric,  was  originally  clothed  in  dry  forest, 
much  of  it  upon  a  series  of  limestone  ter- 
races, the  highest  point  of  which  is  the 
Loma  Gran  Sabana,  having  an  elevation  of 
1082  meters  in  the  north  and  descending  to 
Cerro  Caballo,  and  Loma  de  Chendo,  hav- 
ing elevations  of  322  and  233  meters,  re- 
spectively, to  the  south.  West  of  this  ridge, 
the  land  descends  abruptly  to  Acacia-cac- 
tus desert  to  the  east  of  Cabo  Rojo,  and 
this  habitat  continues  to  the  Dominico- 
Haitian  border  at  Pedernales.  Presumably, 
A.  b.  albocellatus  occurs  throughout  the 
eastern  half  of  the  Peninsula  in  the  for- 
merly high-canopied  forests  of  the  lime- 
stone terraces. 

The  holotype  was  secured  by  Richard 
Thomas  during  the  day  in  a  viny  tangle  in 
semi-xeric  woods  near  Oviedo;  the  lizard 
was  in  an  edge  situation,  since  beyond  the 
dense  vine  tangle  the  woods  thinned  to 
more  scrubby  and  cleared  areas. 

The  name  albocellatus  is  from  the  Latin 


HisPANioLAN  Giant  Anole  •  Schivartz 


119 


"albus"  for  "white"  and  "ocellus"  for  "eye," 
in  allusion  to  the  white  spots  that  are  typi- 
cal of  the  holotype. 

Anolis  baleatus  Cope 

Eti])ristis   baleatus  Cope,    1864,   Pvoc.    Acad.    Nat. 
Sci.    Philadelphia,  p.  168. 

Type  locality.  Santo  Domingo;  holo- 
type, British  Museum  (Natural  History) 
1946.8.29.22. 

Definition.  A  giant  species  of  Hispanio- 
lan  Anolis  characterized  by  the  combina- 
tion of  large  size  (males  to  1<S0  mm, 
females  to  148  mm  snout-vent  length), 
snout  scales  at  level  of  second  canthal 
scales  2  to  5  (modally  2  or  4,  by  popula- 
tion) but  usually  2  or  3  (75  percent),  ver- 
tical loreal  rows  5  to  10  ( modes  by  popula- 
tion 6,  7  or  8),  scales  between  supraorbital 
semicircles  1  to  4  (modally  3),  interpari- 
etal scales  separated  from  supraorbital 
semicircles  modally  by  4  or  5  scales,  verti- 
cal dorsal  scales  generally  small  ( 12  to  24 
in  standard-distance),  ventral  scales  rela- 
tively small  ( 15  to  34  in  standard-distance ) , 
nuchal  crest  scales  in  both  sexes  very  high 
to  high,  rarely  moderate,  never  low,  body 
crest  scales  usually  high  to  moderate,  rarely 
low,  subocular  scales  usually  not  in  contact 
with  supralabial  scales;  dorsal  body  colora- 
tion and  pattern  usually  some  shade  of 
green,  varying  from  dull  greenish  brown  to 
bright  emerald  green,  either  conspicuously 
crossbanded  with  few  (3  or  4)  to  very 
many  crossbands,  in  the  latter  condition  the 
lizards  appearing  tigroid,  or,  on  the  other 
hand,  without  crossbanding  but  blotched, 
never  stiiped  or  with  dark  occipital,  nu- 
chal, or  lateral  dark  markings,  dewlap  in 
males  from  pale  yellow  to  vivid  orange,  in 
females  from  brownish  or  very  pale  yellow 
to  orange  or  gray,  often  suffused  with  gray- 
ish or  brownish,  or  nearly  white,  chin  and 
throat  yellowish,  green,  or  orange,  often 
with  a  dark  dotted  or  mottled  or  reticulate 
pattern,  and  pale  subocular  crescent  absent 
in  adults. 

Distribution.  The  eastern  two-thirds  of 
the    Repiiblica    Dominicana,    from    Puerto 


Plata,  Santiago,  and  La  Vega  provinces 
south  to  San  Cristobal  Province  and  the 
Distrito  Nacional,  and  east  to  La  Altagracia 
Province;  also  in  and  near  the  Sierra  Mar- 
tin Garcia  and  the  southern  slopes  of  the 
Cordillera  Central  and  the  Sierra  de  Ocoa 
in  Azua  and  Peravia  provinces;  occurs  on 
Isla  Saona  but  unrepresented  by  specimens 
from  that  satellite  island. 

Anolis  boleotus  baleatus  Cope 

Type  locality.  "Santo  Domingo";  here 
restricted  to  the  vicinity  of  Puerto  Plata, 
Puerto  Plata  Province,  Repiiblica  Domini- 
cana (see  rationale  for  this  restriction  be- 
low ) . 

Definition.  A  subspecies  of  A.  baleatus 
characterized  by  the  combination  of  mod- 
ally  4  snout  scales  between  second  canthal 
scales,  7  vertical  rows  of  loreal  scales,  3 
scales  between  the  supraorbital  semicircles, 
moderate  number  of  vertical  dorsal  scales 
(14-21;  mean  17.5),  high  number  of  ven- 
tral scales  (19-34;  mean  23.8),  nuchal  crest 
scales  very  high  (usually)  to  high  or  mod- 
erate (rarely),  body  crest  scales  high 
(rarely)  to  moderate  (usually),  subocular 
scales  always  in  contact  with  supralabial 
scales,  males  from  pale  green  or  rich  bluish 
green  to  brown  dorsally,  with  three  bright 
yellow  to  darker  green  or  greenish  brown 
irregular  crossbands,  lower  sides  usually 
bright  yellow,  females  apparently  with  the 
same  body  patterns  and  hues  as  the  males 
(see  below),  throat  in  males  bright  yellow 
to  bright  orange,  rarely  mottled  with 
brown,  dewlap  in  males  always  vivid  to 
brilliant  orange,  and  the  upper  surfaces  of 
hindlimbs  bluish  green,  conspicuously 
barred  with  bright  yellow. 

Distribution.  Known  from  the  Cordil- 
lera Septentrional  and  the  northern  coastal 
plain  of  the  Repiiblica  Dominicana,  from 
Puerto  Plata,  Espaillat,  and  Santiago  prov- 
inces, but  probably  occurring  elsewhere  in 
this  range  and  to  the  north  of  it;  specimens 
from  Los  Bracitos,  Duarte  Province,  should 
also  be  included  ( on  geographical  grounds ) 
with  A.  b.  baleatus,  since  Los  Bracitos  lies 


120         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


in  the  eastern  extremity  of  the  Cordillera  locality  of  the  name  to  the  vicinity  of  a  ma- 
Septentrional,   but  the   specimens   are   old  jor  city  that  is  presumed  to  lie  within  the 
and  greatly  discolored  and  I  have  not  con-  area  to  which  I  ascribe  this  boldly  cross- 
sidered  them  as  pertaining  to  the  nominate  banded  subspecies, 
subspecies.  The  series  of  15  A.  b.  haleatus  shows  the 

Discussion.  Eupristis  haleatus  Cope  was  following  variation.  The  largest  males 
named  from  a  single  specimen  from  "Santo  (ASFS  V33558,  ASFS  V18123)  have  snout- 
Domingo."  I  have  examined  the  holotype,  vent  lengths  of  148,  and  the  largest  fe- 
collected  by  A.  Salle,  in  the  British  Museum  male  ( MCZ  128380)  has  the  same  dimen- 
( Natural  History ) .  Considering  its  length  sion.  These  three  lizards  are  all  from  the 
of  time  in  preservative,  it  is  in  excellent  Cordillera  Septentrional  north  of  Puesto 
condition  and  shows  a  striking  pattern  of  Grande.  Snout  scales  at  level  of  the  second 
three  bold  pale  body  crossbands  on  a  canthal  vary  between  2  and  4;  the  mode 
darker  dorsal  ground  color,  contrastingly  is  4  (eight  specimens).  The  vertical  loreal 
banded  hindlimbs  and  tail,  and  immaculate  rows  vary  between  5  and  9,  with  a  mode 
throat.  The  specimen  is  a  female,  and,  un-  of  7  (six  specimens).  There  are  3  scales 
fortunately,  I  have  only  two  adult  females  between  the  supraorbital  semicircles  in  all 
from  the  range  ascribed  above  to  A.  h.  ha-  specimens.  There  are  modally  5  scales  be- 
leatiis:  both  are  without  color  data  in  life,  tween  the  interparietal  and  the  semicircles; 
At  least  one  of  them  (MCZ  57717)  resem-  5  scales  are  involved  in  63  percent  of  the 
bles  the  pattern  of  the  haleatus  holotype  combination;  actual  counts  are  4/4  (1),  4/5 
to  a  striking  degree.  (4),  5/5  (7),  5/6  (1),  6/6  (1),  and  5/7 

Through  the  courtesy  of  Ernest  E.  Wil-  (1).  Vertical  dorsals  range  between  14  and 
liams,  I  have  a  copy  of  a  map  prepared  by  21  ( mean  17.5 ) ,  horizontal  dorsals  between 
William  J.  Clench  which  shows  the  locali-  16  and  26  (19.7),  and  ventrals  between  19 
ties  where  A.  Salle  is  known  to  have  col-  and  34  (mean  23.8).  Of  nine  adult  males, 
lected.  Considering  the  era  of  his  travels  six  have  the  nuchal  crest  scales  very  high, 
(the  mid-1800's),  Salle  traveled  widely  two  have  these  scales  high,  and  one  has 
throughout  the  Repiiblica  Dominicana,  them  moderate.  Of  three  females,  the  nu- 
from  (in  the  north)  Puerto  Plata,  Ponton,  chal  crest  scales  are  very  high  in  two  and 
Santiago,  Moca,  La  Vega  and  Cotui,  east  high  in  one.  The  body  crest  scales  are  high 
to  Higiiey,  Cabo  Engafio  and  San  Rafael  in  one  male  and  moderate  in  eight  males; 
del  Yuma,  in  the  eastern  interior  to  Hato  in  three  females,  the  body  crest  scales  are 
Mayor  and  El  Seibo,  along  the  southern  high  in  one  and  moderate  in  two.  All  spec- 
coast  from  Santo  Domingo  to  San  Cristo-  imens  have  the  subocular  scales  in  contact 
bal,  Bani,  Azua,  Barreras,  and  Barahona,  with  the  supralabial  scales, 
and  into  the  Valle  de  San  Juan  to  the  city  Males  are  usually  conspicuously  cross- 
of  San  Juan.  He  also  ascended  the  south-  banded.  Specimens  have  been  recorded  as 
ern  slopes  of  the  Cordillera  Central  near  pale  green  with  three  irregular  darker 
San  Jose  de  Ocoa.  Although  much  of  Sal-  green  crossbands,  brown  with  three  faint 
le's  Dominican  travels  was  in  territory  of  green-brown  crossbands,  or  rich  bluish 
A.  haleatus,  he  was  also  in  the  ranges  of  green  with  three  bright  yellow  crossbands. 
A.  ricorcli  and  A.  harahonae.  The  holotype.  The  lower  sides  are  bright  yellow  (which 
as  V^illiams  (1962:  2,  footnote  1)  pointed  grades  into  a  grayish  venter),  and  this  color 
out,  has  elongate  nuchal  crest  scales,  and  also  occurs  on  the  throat,  which  varies 
there  is  no  doubt  that  the  name  haleatus  from  yellowish  to  bright  yellow  or  orange, 
is  applicable  to  some  population  that  pos-  occasionally  mottled  with  brown.  The  dew- 
sesses  this  character.  Since  Salle  traveled  lap  is  brightly  colored;  it  has  been  recorded 
within  the  range  of  the  northern  population  as  "vivid  orange,"  "bright  vivid  orange," 
of  A.  healeatus,  I  have  restricted  the  type  "brilliant  yellow-orange,"  and  "very  bright 


HisPANiOLAN  Giant  Angle  •  Schtvmiz         121 


orange."  The  upper  siirfaee  of  the  head  is 
reddish  brown  and  the  hindhnibs  are  green 
to  bhiish  green,' barred  with  bright  yellow. 
In  general,  male  A.  b.  haleatus  are  vividly 
patterned  and  eolored  lizards.  I  have  eol- 
lected  no  females  myself  and  thus  have  no 
notes  on  this  sex  from  life;  however,  one 
recently  (1971)  collected  female  ( MCZ 
128380)  still  is  dark  green  with  several  thin 
vertical  pale  crossband  remnants  on  the 
sides  and  back,  and  another  female  ( MCZ 
57717)  is  contrastingly  patterned  in  dark 
and  pale  green,  the  latter  occurring  as  ver- 
tical crossbands. 

The  series  includes  three  subadults,  with 
snout-vent  lengths  between  73  and  83. 
One  of  these  (ASFS  V33559;  snout-vent 
length  80)  was  medium  brown  dorsally 
with  a  black  postocular  streak  and  an  or- 
range  dewlap  that  was  streaked  with  black 
basally.  None  of  the  subadults  as  pre- 
served shows  any  crossbanding  or  other 
pattern  elements.  It  is  interesting  that  the 
only  Hispaniolan  giant  anole  taken  at  night 
sleeping  in  the  brown  phase  is  the  above 
mentioned  subadult. 

Remarks.  All  ASFS  specimens  collected 
by  myself  and  parties  were  secured  at 
night  while  the  lizards  were  sleeping. 
Typical  situations  are  in  gallery  forest  and 
cafetales  along  mountain  streams  in  the 
Cordillera  Septentrional.  Favored  sleep- 
ing sites  for  these  lizards  in  the  region  are 
pendant  and  semi-pendant  woody  vines; 
Fowler  reported  that  one  adult  male  se- 
cured by  him  at  night  was  not  asleep  and 
was  slowly  ascending  a  tree  trunk  as 
Fowler  approached.  It  is  possible  that  this 
lizard  had  been  disturbed  by  the  bright 
light  from  Fowler's  flashlight  or  by  unfa- 
miliar movements  and  noises,  since  I  doubt 
that  any  of  the  Hispaniolan  giant  anoles 
are  normally  active  at  night.  However,  all 
these  lizards  waken  quickly  when  dis- 
turbed and  unless  promptly  secured,  grad- 
ually wander  away  into  the  greenery  and 
are  lost  to  view.  One  of  the  juveniles  was 
secured  only  6  feet  (1.8  meters)  above  the 
ground,  whereas  one  of  the  adults  was 
shot  from  a  tree  limb  35  feet  ( 10.7  meters ) 


above  a  mountain  stream.  The  specimen 
from  near  Sosi'ia  was  taken  in  dense  hard- 
woods on  a  limestone  substrate. 

The  altitudinal  distribution  of  A.  1).  ha- 
leatus is  from  1400  to  2200  feet  ( 427  to  671 
meters),  but  the  ta.xon  occurs  much  lower 
than  this,  since  the  specimen  from  near 
Sosua  was  in  limestone  hills  near  sea  level. 

Specimens  examined.  REPUBLICA  DO- 
MINICANA:  Espaillat  Province,  2km  N  Pu- 
esto  Grande,  1400  to  2200  feet  (427  to  671 
meters)  (ASFS  V18048,  ASFS  V33557- 
59);  5  km  N  Puesto  Grande  (MCZ 
128380);  11  km  N  Puesto  Grande,  2100 
feet  (641  meters)  (ASFS  V18123,  ASFS 
V18292):  Puerto  Plata  Province,  11  km  SE 
Sosua  (ASFS  V1717);  Santiago  Province, 
Pena  (MCZ  57713,  MCZ  57715-19);  no  lo- 
cality other  than  Santo  Domingo — British 
Museum  (Natural  History)  1946.9.28.22— 
holotype  of  Eupristis  haleatus. 

Anolis  haleatus  multistruppus 
new  subspecies 

Holotype.  USNM  193975,  an  adult 
male,  from  Guaigiii,  3  mi.  (4.8  km)  S  La 
Vega,  La  Vega  Province,  300  feet  (92  me- 
ters), Repiiblica  Dominicana,  one  of  a  se- 
ries taken  by  Danny  C.  Fowler,  Albert 
Schwartz,  and  Bruce  R.  Sheplan  on  9  No- 
vember 1971.  Original  number  ASFS 
V33680. 

Paratopes.  ASFS  V33681-86,  MCZ 
125612-15,  CM  54107-12,  same  data  as  ho- 
lotype; ASFS  V18547-50,  same  locality  as 
holotype,  J.  R.  Dennis,  J.  A.  Rodgers,  Jr., 
and  A.  Schwartz,  27  July  1969. 

Definition.  A  subspecies  of  A.  haleatus 
characterized  by  the  combination  of  mod- 
ally  2  snout  scales  between  second  canthal 
scales,  7  vertical  rows  of  loreal  scales,  3 
scales  between  the  supraorbital  semicircles, 
4/4  scales  between  the  interparietal  and  the 
supraorbital  semicircles,  high  number  of 
vertical  dorsal  scales  (14-24;  mean  18.6), 
moderate  number  of  ventral  scales  ( 18-29; 
mean  22.3),  nuchal  crest  scales  very  high 
to  high  (usually)  to  moderate  (rarely), 
body  crest  scales  high  (rarely)  to  moder- 
ate  (usually),  subocular  scales  almost  al- 


122         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


ways  separated  from  supralabial  scales  by 
one  row  of  scales,  both  sexes  as  adults  re- 
taining the  complex  juvenile  pattern  of 
many  fine  dark  green,  green,  and  yellow 
vertical  bars,  occasionally  (in  females) 
bright  pea  green  with  three  pale  green 
crossbars  more  prominent  than  any  other 
dorsal  pattern  elements,  throat  green  to 
yellow  green,  dewlap  in  males  very  pale 
yellow  to  very  pale  peach,  suffused  basally 
with  pale  gray,  in  females  very  pale  yellow 
to  pale  yellow,  strongly  suffused  with  pale 
gray  to  entirely  pale  gray. 

Distribution.  Known  only  from  the  type 
locality  but  presumed  to  occur  on  the 
northern  and  probably  eastern  lower  faces 
of  the  Cordillera  Central  in  proper  habi- 
tats; possibly  extending  as  far  west  on  the 
northern  face  of  this  range  as  the  Rio  Bao 
near  Los  Montones  (see  discussion  below). 

Description  of  holotype.  An  adult  male 
with  a  snout-vent  length  of  146  and  a 
tail  length  (broken)  of  97;  snout  scales  at 
level  of  second  canthals  2,  9  vertical  rows 
of  loreal  scales,  3  scales  between  the  su- 
praorbital semicircles,  4/4  scales  between 
the  interparietal  and  the  supraorbital  semi- 
circles, vertical  dorsals  21,  horizontal  dor- 
sals 25,  ventrals  29,  one  row  of  scales  be- 
tween the  suboculars  and  supralabials, 
fourth  toe  lamellae  on  phalanges  II  and  III 
30,  nuchal  crest  scales  high,  body  crest 
scales  moderate;  in  life,  dorsal  body  pat- 
tern of  many  fine  green,  dark  green,  and 
yellow  crossbands,  upper  surface  of  head 
grayish  tan  in  contrast  to  the  brighter  dor- 
sal colors,  chin  and  throat  very  pale  yellow 
or  yellow-green  with  no  clearly  delineated 
darker  green  markings,  and  dewlap  very 
pale  yellow,  much  suffused  basally  with 
gray. 

Variation.  The  series  of  21  A.  h.  multi- 
struppus  is  composed  of  eight  males  and  13 
females.  The  largest  male  has  a  snout-vent 
length  of  146  and  is  the  holotype.  The 
largest  female  (ASFS  V33684)  has  a 
snout-vent  length  of  136  and  is  a  topo- 
type.  Snout  scales  at  the  level  of  the  sec- 
ond canthals  range  between  2  and  5;  the 
mode   is   2    (15  specimens).    The  vertical 


loreal  rows  vary  between  6  and  9,  with  a 
mode  of  7  (nine  specimens).  There  are  2 
or  3  scales  between  the  supraorbital  semi- 
circles (mode  3).  There  are  modally  4 
scales  between  the  interparietal  and  the 
supraorbital  semicircles;  4  scales  are  in- 
volved in  58  percent  of  the  combinations; 
actual  counts  are  3/3  (1),  3/4  (2),  4/4 
(8),  4/5  (5),  5/5  (3),  and  5/6  (1).  Ver- 
tical dorsals  range  between  14  and  24 
(mean  18.6),  horizontal  dorsals  between 
17  and  25  (20.6),  and  ventrals  between  18 
and  29  (22.3).  All  three  adult  males  have 
the  nuchal  crest  scales  high,  and  of  ten 
females,  three  have  these  scales  very  high, 
five  have  them  high,  and  two  have  them 
moderate.  All  three  males  have  the  body 
crest  scales  moderate,  whereas  two  females 
have  the  body  crest  scales  high,  eight  have 
them  moderate,  and  one  has  them  low.  In 
all  but  one  specimen  (5  percent),  the  sub- 
oculars  are  separated  from  the  supralabials 
by  1  scale. 

Adults  of  both  sexes  retain  the  juvenile 
multibanded  pattern  of  dark  greens,  me- 
dium greens,  and  yellow.  One  adult  fe- 
male was  recorded  as  bright  pea-green 
with  three  pale  green  crossbands,  which 
are  remnants  of  the  hollowed  yellow  cen- 
ters of  the  five  or  six  dark  brown  to  dark 
green  crossbands.  In  general,  the  total  as- 
pect of  adults  and  juveniles  is  of  a  con- 
trastingly tigroid  lizard,  the  stripes  varying 
shades  of  greens,  yellows,  and  ( in  the  dark 
phase)  browns.  The  upper  surface  of  the 
head  is  grayish  tan  in  males  and  tannish 
green  in  females,  and  the  throat  is  un- 
marked green  to  yellow-green.  One  of  the 
most  striking  features  of  A.  b.  multistrup- 
pus  is  the  faded  dewlap  coloration.  In 
males,  the  colors  vary  between  very  pale 
yellow  and  very  pale  peach,  basally  suf- 
fused with  pale  gray.  In  females,  the  dew- 
lap is  even  more  drab,  with  pale  yellow 
the  basic  color,  but  the  gray  suffusion  may 
be  so  extensive  as  to  limit  the  yellow  pig- 
ment to  the  dewlap  edge  or  to  cause  the 
dewlap  to  be  pale  gray. 

The  type  series  includes  seven  juveniles 
and  subadults,  with  snout-vent  lengths  be- 


HisPANioLAN  Giant  Angle  •  Schwartz 


123 


twecn  47  and  99.  These  prcstMit  a  imi- 
fonii  aspect  of  multiple  dorsal  bands  as 
described  above,  and  even  tlie  largest  of 
the  subadnlts  clearly  shows  this  condition. 
In  life,  a  small  juvenile  (snout-vent  length 
53)  was  recorded  as  pale  gray  with  a 
yellow-green  head  and  about  four  reversed 
chevrons  between  the  neck  and  the  hind- 
limbs,  these  chevrons  being  the  pale  hol- 
lowed remnants  of  the  darker  crossbands, 
wliich,  in  this  individual,  arc  obscure.  The 
small  lizard  also  had  a  black  postocular 
line  and  a  charcoal  postangular  smudge. 
The  juvenile  and  subadult  dewlaps  are 
pale  flesh  to  very  pale  yellow,  somewhat 
suffused  basally  with  light  to  very  dark 
gray. 

Comparisom.  Meristically,  inulfistrup- 
pus  differs  from  nominate  haleatus  in  hav- 
ing 2  (rather  than  4)  snout  scales  at  the 
level  of  the  second  canthals,  4/4  (rather 
than  5/5)  scales  between  the  interparietal 
and  the  supraorbital  semicircles,  and  in 
having  slightly  less  ventrals  (means  22.3 
and  23.8).  There  is  also  a  strong  tendency 
for  both  sexes  of  haleatus  to  have  very 
high  nuchal  crest  scales,  whereas  these 
scales  are  more  often  only  high  in  midti- 
struppus.  It  is  in  color  and  pattern  that 
these  two  subspecies  differ  most  strikingly. 
In  the  introduction  to  the  present  paper  I 
commented  on  my  having  collected  speci- 
mens from  the  Cordillera  Septentrional  and 
Guaigiii  on  two  succeeding  days,  and  on 
the  color  and  pattern  differences  being  at 
once  very  apparent.  The  bright  orange 
throat  and  dewlap  of  haleatus  contrast 
quite  obviously  with  the  pale  yellow  to 
gray  dewlaps  in  multistruppus.  The  body 
patterns  of  the  two  subspecies  likewise  are 
quite  different,  with  the  finely  and  multi- 
banded  multistruppus  in  contrast  to  the  ir- 
regularly banded  dorsum  with  only  three 
bands  in  ])aleatus. 

Discussion.  A.  h.  multistruppus  is 
known  with  certainty  from  only  a  single 
locality,  which  lies  at  the  foot  of  the  Cor- 
dillera Central  at  an  elevation  of  300  feet 
(92  meters).  The  locality  is  unique  in  that 
it  represents  an  extensive  stand  of  original 


lowland  forest  in  this  region,  hardwood 
forest  which  abuts  upon  the  lower  pine- 
clad  slopes  of  the  mountains.  This  locality, 
Guaigiii,  is  separated  from  the  known 
range  of  A.  h.  haleatus  by  the  Vallc  de 
Cibao,  which  here  is  a  moderately  arid 
and  broad  valley  presently  very  much  un- 
der cultivation.  I  have  seen  no  specimens 
from  this  intervening  valley  but  surely  the 
lizards  occur  there,  despite  the  cultivation. 

One  other  specimen  requires  mention. 
This  is  a  subadult  male  (ASFS  V33856) 
with  a  snout-vent  length  of  55,  from  3.4 
mi.  (5.4  km)  SE  Los  Montones,  Rio  Bao, 
1600  feet  (488  meters).  This  locality  is  on 
the  northern  slopes  of  the  Cordillera  Cen- 
tral, some  45  kilometers  to  the  west  of 
Guaigiii,  but  separated  from  Guaigiii  by 
intervening,  moderately  high  spurs  of  the 
Cordillera  Central.  The  specimen  was  se- 
cured by  a  local  boy  in  an  area  of  high- 
canopied  forest  along  the  Rio  Bao.  A  visit 
by  ourselves  to  this  area  at  night  yielded 
no  A.  haleatus,  despite  exceptionally  fine 
conditions.  The  lizard  in  life  was  all  green 
except  for  a  white  preaxillary  bar,  and  the 
dewlap  was  dull  brownish.  This  specimen 
in  no  way  resembles  comparably  sized  ju- 
venile multistruppus,  in  either  color  or  pat- 
tern.   Its  status  remains  uncertain. 

To  the  east,  multistruppus  must  come  in 
close  contact  or  intergrade  with  the  sub- 
species that  occurs  throughout  the  north- 
eastern portion  of  the  Republica  Domini- 
cana;  details  of  this  contact  will  be 
discussed  under  the  account  of  the  latter 
subspecies.  Likewise,  to  the  south,  multi- 
struppus may  come  into  contact  with  the 
subspecies  in  the  high  Cordillera  Central; 
details  of  this  association  will  be  discussed 
under  the  description  of  the  Central  sub- 
species. 

Remarks.  All  specimens  of  A.  h.  multi- 
struppus were  collected  on  two  occasions, 
while  the  lizards  slept  at  night.  Young  in- 
dividuals were  taken  from  generally  low 
situations  on  shrubs  and  the  lower 
branches  of  trees,  whereas  adults  were  ob- 
served sleeping  in  the  higher  canopy;  the 
total  range  of  heights  was  between  5  feet 


124         Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  2 


and  25  feet  (1.5  and  7.6  meters).   The  Rio  scales    always    high,    subocular    scales    al- 

Camu  flows  through  the  Guaigiii  woods,  most    always    separated    from    supralabial 

and    many    individuals    were    taken    from  scales   by   one   row  of  scales,   both   sexes 

tree  limbs  that  overhang  the  river.  either  marbled  or  blotched  with  varying 

The    name    multistruppus    is    from    the  shades  of  greens  or  browns,  or  dark  brown 

Latin  "multus"  for  "many"  and  "struppus"  banded  with  dull  cream,  never  with  many 

for  "thong,  strap,"  in  allusion  to  the  many  fine  crossbars,  venter  in  males  pale  green, 

dorsal  crossbands  in  this  subspecies.  flecked  with  darker  green,  and  male  dew- 
laps  pale   yellow-orange   to   orange,    gray 

Anolis  baleatus  sublimis  new  subspecies  basally  and  marbled  green  anteriorly,  fe- 

Holotijpe.     CM    54104,    an    adult   male,  male    dewlaps     irregularly    yellow-orange 

from  0.3  mi.  (0.5  km)  E  El  Rio,  3800  feet  with  brown  spotting. 

(1159  meters).  La  Vega  Province,  Repub-  Distribution.     The  uplands   of  the   Do- 

lica  Dominicana,  taken  by  Richard  Thomas  minican    Cordillera    Central    at   elevations 

on  26  June  1963.    Original  number  ASPS  between  2000  and  4000  feet,  in  the  area 

X8114.  between  El  Rio,  La  Palma,  and  Jarabacoa. 

Paratypes.     ASFS  X8558,  4  km  SW  El  Description  of  holotype.     An  adult  male 

Rio,    4000   feet    ( 1220   meters ) ,    La   Vega  with  a  snout-vent  length  of  143  and  a  tail 

Province,    Republica    Dominicana,    R.    F.  length  of  167  (regenerated);  snout  scales 

Klinikowski,  2  July  1963;  USNM  62104-05,  at  level  of  second   canthals  3,   7  vertical 

El  Rio,  La  Vega  Province,  Republica  Do-  rows  of  loreal  scales,  2  scales  between  the 

minicana,    W.    L.    Abbott,    19    May    1919;  supraorbital    semicircles,    4/4    scales    be- 

AMNH  41294,  El  Rio,  La  Vega  Province,  tween   the  interparietal   and  the  supraor- 

Repiiblica   Dominicana,   G.    K.    Noble,   31  bital  semicircles,  vertical  dorsals  20,  hori- 

August  1922;  ASFS  V18594,  La  Palma,  14  zontal  dorsals  21,  ventrals  29,  one  row  of 

km  E  El  Rio,  3500  feet  (1068  meters).  Re-  scales  between  the  suboculars  and  supra- 

publica  Dominicana,  J.  A.  Rodgers,  Jr.,  30  labials,  fourth  toe  lamellae  on  phalanges  II 

July  1969;  MCZ  107019-21,  La  Palma,  14  and  III  31,  nuchal  crest  scales  and  dorsal 

km  E  El  Rio,  3500  feet  ( 1068  meters ) ,  La  body    crest    scales    high;    in    life,    dorsum 

Vega  Province,  Republica  Dominicana,  na-  dark  brown  banded  with  dull  cream,  this 

tive  collectors  for  E.  E.  Williams  and  A.  S.  pattern    extending    onto    the   tail,    eyeskin 

Rand,  25-31  July  1968;   MCZ  128397,  La  grayish  with  a  pale  yellow  eyering,  venter 

Palma,  14  km  E  El  Rio,  3500  feet  ( 1068  pale  green,  flecked  with  darker  green,  chin 

meters).  La  Vega  Province,  Republica  Do-  and   throat   concolor   with   and   patterned 

minicana,  T.  P.  Webster  and  R.  B.  Huey,  like    venter,    dewlap    pale    yellow-orange, 

6  July   1971;   ASFS  V18363-69,   8  km  W  grayish  basally  and  marbled  with  green  an- 

Jarabacoa,    2000    feet    (610    meters).    La  teriorly. 

Vega  Province,  Republica  Dominicana,  J.  Variation.     The  series  of  18  sublimis  is 

A.  Rodgers,  Jr.,  19  July  1969.  composed  of  nine  males  and  nine  females. 

Definition.     A  subspecies  of  A.  baleatus  The  largest  male    (USNM   62104)    has   a 

characterized  by  the  combination  of  mod-  snout-vent    length    of     150,     the     largest 

ally  2  snout  scales  between  second  canthal  female    (MCZ   107021)    141;   the   male   is 

scales,  7  vertical  rows  of  loreal  scales,  3  from  El  Rio,  the  female  from  La  Palma. 

scales    between    the    supraorbital    semicir-  Snout   scales   at   the   level   of   the   second 

cles,  4/4  scales  between  the  interparietal  canthals  range  between  2  and  5;  the  mode 

and  the  supraorbital  semicircles,  high  num-  is  2   (10  specimens ) .    The  vertical  loreal 

ber  of  vertical  dorsal  scales  (17-21;  mean  rows  vary  between  6  and  9,  with  a  mode  of 

19.2 ) ,  high  number  of  ventral  scales  ( 19-  7    ( eight   specimens ) .    There   are   2   to   4 

32;  mean  25.1),  nuchal  crest  scales  very  scales    between   the    supraorbital    semicir- 

high  (usually)  to  high  (rarely),  body  crest  cles  (mode  3).   There  are  modally  4  scales 


HisPANioLAN  Giant  Angle  •  Schwartz         125 


between  the  interparietal  and  the  supraor- 
bital semieircles;  4  scales  are  involved  in 
65  percent  of  the  combination;  the  actual 
counts  are  3/3  ( 1),  3/4  ( 1),  4/4  ( 10),  4/5 
(1),  5/5  (3),  and  5/6  (1).  Vertical  dor- 
sals range  between  17  and  21  (mean  19.2), 
horizontal  dorsals  between  17  and  24 
(20.4),  and  ventrals  between  19  and  32 
(25.1).  Of  the  six  adult  males,  three  have 
the  nuchal  crest  scales  very  high  and  three 
ha\e  them  high,  whereas  all  five  adult  fe- 
males have  these  scales  very  high.  All 
adults  of  both  sexes  have  the  dorsal  body 
crest  scales  high.  Three  lizards  ( 17  per- 
cent) have  the  subocular  scales  in  contact 
with  the  supralabials. 

In  the  green  phase,  adults  of  both  sexes 
are  irregularly  marbled  or  blotched  with 
varying  shades  of  green  or  browns, 
whereas  in  the  brown  phase,  the  body  is 
dark  brown  with  three  cream  crossbands. 
In  males  the  venter  and  the  chin  and 
throat  are  pale  green,  flecked  or  mottled 
with  darker  green,  the  flecking  or  mottling 
variably  expressed  in  the  series.  The  dew- 
lap in  males  is  pale  yellow-orange  to  or- 
ange, gray  basally  and  often  with  marbled 
green  markings  anteriorly,  these  markings 
being  a  continuation  of  the  dark  green 
throat  markings.  In  females,  the  dewlap  is 
irregularly  mottled  with  yellow-orange  and 
has  some  brown  spotting.  As  preserved, 
the  series  is  remarkably  uniform  in  show- 
ing vague  pale-and-dark  marblings  or  mot- 
tlings,  and  no  adult  shows  any  indication 
of  crossbands. 

The  series  of  paratypes  includes  six  ju- 
veniles and  subadults,  with  snout-vent 
lengths  between  49  and  94.  The  three 
smallest  of  these  (49-70)  are  presently 
patternless,  as  is  also  a  specimen  with  a 
snout-vent  length  of  73.  Two  other  sub- 
adults  (snout-vent  lengths  75  and  94) 
show  vague  indications  of  mottled  dorsum 
with  (in  the  larger)  three  slightly  paler 
dorsal  crossbands.  The  larger  of  these  two 
specimens  was  recorded  in  life  as  dark 
green  dorsally  with  pale  green  crossbands, 
and  the  interbars  are  mottled  or  marbled 
with  greens.    The  smallest  juvenile  noted 


above  was  bright  yellow-green  in  life  and 
had  the  venter  slightly  paler  yellow-green; 
the  concealed  surfaces  of  the  thighs  were 
lead-gray,  bordered  above  by  bufl^y.  The 
absence  of  pale  crossbars  in  very  young 
specimens  of  S'tihli)nis  is  noteworthy 

Comparisons.  There  are  no  meristic 
counts  that  separate  sul)Iimis  from  adja- 
cent imiltisfruppus;  the  means  of  ventral 
scales  in  the  two  subspecies  difl^er  slightly, 
however  (22.3  in  sul)li77iis,  25.1  in  multi- 
struppus).  There  is  also  a  tendency  for 
su])limis  to  have  more  consistently  very 
high  to  high  nuchal  crest  scales.  The  two 
subspecies  diff^er  abundantly  in  body  pat- 
tern, however,  with  miilfistruppus  having 
many  fine  dorsal  crossbands  and  sublimis 
having  basically  a  blotched  dorsal  pattern 
with  three  bars  present  in  some  instances. 
The  juveniles  of  these  two  subspecies  are 
equally  as  distinct  as  the  adults  are  in  dor- 
sal body  pattern.  The  dewlaps  are  brighter 
in  male  su])limis  than  in  male  muUistnip- 
pus,  the  latter  tending  toward  pale  yellow 
and  yellow-grays,  whereas  in  the  former 
the  dewlaps  are  yellow-orange  to  orange, 
although  there  is  a  gray  basal  suff^usion. 
The  ventral  and  throat  flecking  or  mot- 
tling in  siihlimis  differs  from  the  unmarked 
condition  in  mtiltistruppiis. 

A.  h.  sublimis  differs  from  A.  h.  hcileatiis 
in  having  2  (rather  than  4)  snout  scales  at 
the  level  of  the  second  canthals,  4/4 
(rather  than  5/5)  scales  between  the  in- 
terparietal and  supraorbital  semicircles, 
higher  means  of  vertical  dorsal  scales  ( 19.2 
versus  17.5)  and  ventrals  (25.1  versus 
23.8).  Both  subspecies  have  very  high  to 
high  nuchal  crest  scales.  In  color,  haleaius 
is  much  the  brighter,  with  an  immaculate 
bright  yellow  to  orange  throat  and  bright 
yellow  to  orange  dewlap  in  males,  whereas 
the  dewlaps  in  sublimis  are  as  bright  as 
those  in  haleatus  but  have  a  gray  basal 
wash.  The  patterned  throat  and  venter  in 
sublimis  differ  from  the  immaculate  throat 
in  buleatus.  The  dorsal  patterns  of  these 
two  subspecies  likewise  are  quite  different, 
that    of    baleatus    regularly    consisting    of 


126         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


three  pale  crossbands,  whereas  that  of  sub- 
limis  is  mottled  or  blotched. 

Discussion.  A.  h.  suhlimis  is  closest 
geographically  to  miiltistruppus;  the  two 
subspecies  are  known  from  localities  sep- 
arated by  only  20  kilometers  airline 
(Guaigi^ii  and  8  km  W  Jarabacoa),  but 
minimally  by  a  1700-foot  (519  meters)  dif- 
ference in  elevation  and  by  extensive 
stands  of  pine  forest,  a  habitat  which  no 
Hispaniolan  giant  anole  occupies.  All  spec- 
imens of  suhlimis  were  collected  in  mon- 
tane gallery  forest  along  streams,  and  the 
subspecies  appears  to  be  restricted  to  this 
sort  of  situation.  Rand  and  Williams 
(1969:  9)  noted  that  they  collected  one 
juvenile  about  10  feet  (3  meters)  up  on  a 
small  branch  of  a  forest  tree  at  La  Palma, 
and  that  two  adults  were  brought  to  them 
by  natives  from  a  large  tree  in  a  nearby 
agricultural  area.  A.  h.  suhlimis  is  thus 
not  known  to  come  into  contact  with  multi- 
struppus  on  the  northern  slopes  of  the  Cor- 
dillera Central  nor  with  the  yet-to-be-de- 
scribed subspecies  to  the  east  in  the 
Dominican  lowlands.  Likewise,  it  should 
be  recalled  that  the  southwestern  slopes  of 
this  range  are  occupied  by  A.  r.  ricordi; 
the  nearest  localities  for  ricordi  and  sul)- 
lirnis  (Juan  de  Herrera;  south  of  El  Rio) 
are  separated  by  about  70  kilometers  air- 
line, but  this  intei^vening  area  is  composed 
of  the  rugged  and  very  high  massif  of  the 
Cordillera  Central  whose  upper  elevations 
are  covered  by  pine.  It  seems  unlikely  that 
ricordi  and  suhlimis  come  into  contact  di- 
rectly across  the  Cordillera. 

The  juvenile  (ASFS  33856)  from  near 
Los  Montones  upon  which  I  commented 
in  the  discussion  of  A.  h.  miiltistruppus 
may  be  correctly  assigned  to  suhlimis,  since 
the  habitat  and  elevation  for  that  specimen 
is  much  more  like  that  for  suhlimis  than 
multistruppus.  In  color  and  lack  of  pattern 
it  agrees  quite  well  with  small  suhlimis, 
but  until  adults  have  been  collected  in  the 
Los  Montones  region  (which  lies  some  30 
kilometers  to  the  northwest  of  Jarabacoa, 
the  closest  suhlim.is  locality)  I  am  reluc- 
tant to  extend  the  known  range  of  suhlim,is 


into  that  area.  It  is  this  Los  Montones  A. 
haleatus  which  is  closest  geographically 
(50  kilometers)  to  an  A.  ricordi  locality 
( Los  Quemados )  in  the  northwestern  por- 
tion of  the  Republica  Dominicana. 

Remarks.  All  ASFS  A.  h.  suhlimis  were 
taken  at  night  while  asleep.  All  situations, 
as  noted  above,  were  stream-associated 
hardwood  forest  and  cafetales,  and  the  liz- 
ards slept  on  vines  and  branches  in  their 
customary  fashion.  The  restriction  of  suh- 
limis to  riverine  gallery  forest  is  doubtless 
artificial,  since  it  is  only  along  rivers  and 
streams  in  this  area  that  any  of  the  original 
montane  hardwood  forests  still  remain.  In 
one  case  (west  of  Jarabacoa)  the  stream 
was  extremely  steep,  whereas  in  others  the 
streams  were  level.  The  altitudinal  distri- 
bution ( to  which  the  name  suhlimis  refers ) 
is  high.  Only  A.  r.  viculus  reaches  as  high 
an  elevation  in  the  Massif  de  la  Hotte  in 
southwestern  Haiti. 

Anolis  baleofus  caeruleolatus 
new  subspecies 

Holotype.  USNM  193976,  an  adult 
male,  from  1.0  mi.  (1.6  km)  S  Caiio  Abajo, 
>/Iaria  Trinidad  Sanchez  Province,  Repub- 
lica Dominicana,  one  of  a  series  collected 
by  native  collectors  on  28  November  1971. 
Original  number  ASFS  V34486. 

Paratypes.  CM  54119-26,  MCZ  125628- 
33,  ASFS  V34502-13,  same  data  as  holo- 
type; AMNH  6017,  Villa  Riva,  Duarte 
Province,  Republica  Dominicana,  C.  R. 
Halter,  May-July  1915. 

Associated  specimens.  REPCBLICA 
DOMINICANA:  Duarte  Province,  Los 
Bracitos  (AMNH  41465-66);  ca.  4  km  NE 
Ponton  (Rio  Cuaba)  (ASFS  V2987);  San- 
chez Ramirez  Province,  1  km  SE  La  Mata 
(ASFS  V33650-51);  La  Vega  Province, 
12.8  km  NW  Bonao,  1200  feet  (366  meters) 
(ASFS  V4317);  71  km  NW  Santo  Domingo 
(=  near  La  Cumbre)  (MCZ  128369);  San 
Cristohal  Province,  5.0  mi.  (8.0  km)  NE 
Gonzalo,  1000  feet  (305  meters)  (ASFS 
V29420-21). 

Definition.  A  subspecies  of  A.  haleatus 
characterized  by  the  combination  of  mod- 


HisPANioLAN  Giant  Anole  •  Schwartz 


127 


ally  4  scales  between  second  canthal  scales, 
8  vertical  rows  of  loreal  scales,  3  scales  be- 
tween the  supraorbital  semicircles,  5/5 
scales  between  the  interparietal  and  the 
supraorbital  semicircles,  moderate  number 
of  vertical  dorsal  scales  (14-22;  mean 
17.1),  moderate  number  of  ventral  scales 
(15-32;  mean  22.4),  nuchal  crest  scales 
very  high  to  high  (usually)  to  moderate 
or  even  low  (rarely)  in  both  sexes,  body 
crest  scales  extremely  variable,  modally 
moderate  in  both  sexes,  but  with  some  oc- 
cinrences  of  high  and  many  occurrences 
of  low  body  crest  scales,  subocular  scales 
almost  always  separated  from  supralabial 
scales  by  one  row  of  scales,  both  sexes 
some  shade  of  green  (usually  dark)  with 
foiu-  pale  green  crossbars  and  with  bright 
sky-bhie  blotches  along  the  junction  of  the 
green  dorsal  color  and  the  paler  venter 
(less  prominent  in  females  than  in  males), 
dewlap  in  males  pale  yellow  to  orange,  in 
females  pale  yellow  to  orange  but  with 
much  dark  brown  to  grayish  streaking  or 
smudging,  throat  in  males  deep  yellow-or- 
ange and  immaculate  or  with  very  faint 
greenish  dots,  in  females  yellow-green  to 
bright  yellow,  always  with  some  darker 
green  dots,  rarely  marbled  with  dark  green, 
but  never  streaked  with  that  color. 

Distribution.  Northeastern  Republica 
Dominicana,  from  Duarte,  Sanchez  Rami- 
rez, La  Vega,  and  northern  and  eastern 
San  Cristobal  provinces,  to  the  base  of  the 
Peninsula  de  Samana  (Caiio  Abajo);  in- 
tergrades  with  the  subspecies  to  the  south 
and  east  in  the  region  of  El  Seibo  Province. 

Description  of  holotijpe.  An  adult  male 
with  a  snout-vent  length  of  137  and  a  tail 
length  of  250;  snout  scales  between  second 
canthals  4,  7  vertical  rows  of  loreal  scales, 
3  scales  between  the  supraorbital  semicir- 
cles, 6/6  scales  between  the  interparietal 
and  supraorbital  semicircles,  vertical  dor- 
sals 16,  horizontal  dorsals  23,  ventrals  26, 
one  row  of  scales  between  the  suboculars 
and  supralabials,  fourth  toe  lamellae  on 
phalanges  II  and  III  30,  nuchal  crest  scales 
very  high,  body  crest  scales  moderate;  in 
life,    dorsum    dark    green    with    four    pale 


green  crossbars,  the  dark  green  color 
blending  fjuickly  at  the  junction  of  the  dor- 
sal and  ventral  color  into  a  series  of  diag- 
onally directed  sky-blue  areas  that  give  a 
ragged  appearance  to  the  jmiction  of  the 
dorsal  and  ventral  colors;  dorsal  crossbands 
continue  onto  the  tail;  cascjue  gray-green, 
eyeskin  pale  pea-green;  dewlap  pale  yel- 
low-orange, chin  slightly  deeper  yellow-or- 
ange, throat  yellow-orange,  immaculate 
except  for  some  vague  pale  greenish 
smudges  posterolaterally. 

Variation.  The  series  of  A.  b.  caeruleo- 
lattis  consists  of  20  males  and  17  females. 
The  largest  male  (ASFS  V34505)  has  a 
snout-vent  length  of  148,  the  largest  fe- 
male (AMNh'  6017)  145.  The  male  is 
from  the  type  locality,  the  female  from 
Villa  Riva.  Snout  scales  at  the  level  of  the 
second  canthals  range  between  2  and  5; 
the  mode  is  4  ( 14  specimens ) .  The  verti- 
cal loreal  rows  vary  between  6  and  10;  the 
mode  is  8  ( 15  specimens ) .  There  are  2  or 
3  scales  between  the  supraorbital  semicir- 
cles (mode  3).  There  are  modally  5  scales 
between  the  interparietal  and  the  supraor- 
bital semicircles;  5  scales  are  involved  in 
52  percent  of  the  combinations;  actual 
counts  are  4/4  (3),  4/5  (6),  5/5  (10),  5/6 
(7),  6/6  (4),  6/7  (1),  4/6  (1),  and  5/7 
( 1 ) .  Vertical  dorsals  range  between  14 
and  22  (mean  17.1),  horizontal  dorsals  be- 
tween 15  and  25  ( 19.9 ) ,  and  ventrals  be- 
tween 15  and  32  ( 22.4 ) .  Of  16  adult  males, 
four  have  the  nuchal  crest  scales  very  high, 
11  have  them  high,  and  one  has  them  mod- 
erate. Of  17  adult  females,  four  have  the 
nuchal  crest  scales  very  high,  ten  have 
them  high,  and  three  have  them  moderate. 
In  the  adult  males,  the  body  crest  scales 
are  high  in  six  males,  moderate  in  eight, 
and  low  in  two,  whereas  in  the  adult  fe- 
males, these  scales  are  high  in  five,  mod- 
erate in  six,  and  low  in  six.  All  but  two 
lizards  (6  percent)  have  the  suboculars 
separated  by  one  row  of  scales  from  the 
supralabials. 

In  a  series  of  12  adult  male  topotypes, 
the  dorsal  ground  color  was  recorded  as 
some  shade  of  green  (usually  dark  green) 


128         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


with  four  pale  pea-green  crossbands.  The  patterned  hke  adults  except  that  the  sky- 
dorsal  green  color  blends  quickly  ventro-  blue  lower  edges  to  the  dorsal  color  were 
laterally  into  a  series  of  irregular  sky-blue  absent  and  the  dewlap  was  streaked  brown 
patches  or  blotches  that  mark  the  border  and  gray  basally.  The  chin  and  throat 
between  the  dorsal  green  and  the  pale  yel-  were  immaculate  pale  green.  There  are  no 
low  to  cream  venter.  These  sky-blue  color  data  on  the  other  juveniles,  and  none 
patches  are  often  prominently  extended  of  them  presently  shows  any  pattern, 
onto  the  lateral  margins  of  the  venter  as  a  Comparisons.  A.  h.  caeruleolatus  dif- 
series  of  diagonal,  posteriorly  directed  fers  from  all  previously  described  subspe- 
areas,  which,  upon  preservation,  are  still  cies  in».having  the  sky-blug  patching  along 
prominent  features  of  the  lower  sides.  The  the  lower  sides.  In  having  four  dorsal  pale 
upper  surface  of  the  head  was  gray-green  green  body  bands,  caeruleolatus  differs 
to  brown,  the  eyeskin  pale  pea-green.  The  strikingly  from  multistruppus  with  its  mul- 
dorsal  banded  pattern  of  dark  and  light  tiple  banding;  in  addition,  the  dewlap  of 
green  continues  onto  the  tail.  The  dewlap  multistruppus  is  pale  and  often  grayish,  in 
is  pale  yellow-orange,  yellow,  or  orange,  contrast  to  the  generally  brighter  dewlaps 
and  the  chin  is  slightly  deeper  yellow-or-  of  caeruleolatus.  From  nominate  baleatus, 
ange,  concolor  with  the  throat,  which  is  caeruleolatus  differs  in  having  the  throat 
either  immaculate  (usually)  or  with  very  yellow  to  yellow-green  rather  than  bright 
faint  greenish  dots  or  smudges.  Eleven  fe-  yellow  to  orange,  and  female  caeruleolatus 
male  topotypes  were  colored  and  patterned  have  the  throat  with  dark  green  markings, 
dorsally  like  the  males,  with  the  pattern  From  high  upland  sublitnis,  caeruleolatus 
extending  onto  the  tail,  but  there  is  only  a  differs  in  having  the  sky-blue  blotches  ven- 
vague  indication  of  the  ventrolateral  sky-  trolaterally  and  in  lacking  ventral  mark- 
blue  pigmentation.  The  necks  of  females  ings,  and  whereas  caeruleolatus  has  com- 
were  often  streaked  with  dark  and  pale  parably  pigmented  dewlaps^  those  in 
greens.  The  chin  and  throat  were  yellow  sublimis  are  generally  paler  and  often  suf- 
to  yellow-green,  regularly  with  some  fus^d  at  least  basally  with  gray.  The  dor- 
darker  green  dots,  blotches,  or  occasionally  sal  patterns  of  both  sublimis  and  caeruleo- 
marbled  with  dark  green.  The  female  dew-  latus  are  comparable,  since  both  are 
lap  was  yellow  to  pale  orange,  streaked  crossbanded. 
with  dark  brown  or  grayish.  As  far  as  meristic  counts  are  concerned, 

Two  females  from  the  haitises  region  caeruleolatus  differs  from  the  named  sub- 
near  Gonzalo  were  deep  to  emerald  green  species  in  the  following  ways.  Compared 
in  life  with  yellow  dewlaps  having  varying  with  baleatus,  caeruleolatus  has  modally  8 
amounts  of  brown  streaking  or  smudging;  (rather  than  7)  vertical  loreal  rows,  and  a 
the  limbs  were  contrastingly  banded  dark  lower  mean  number  of  ventral  scales  (22.4 
and  pale  green.  The  throats  were  bright  versus  23.8).  There  is  also  a  strong  ten- 
yellow  to  bright  green,  with  scattered  dency  for  adult  caeruleolatus  to  have  mod- 
deeper  green  spots  in  each  case.  In  a  pair  erate  to  low  body  crest  scales,  whereas  in 
from  La  Mata,  the  dorsa  were  bright  baleatus  the  tendency  is  toward  high  to 
green,  somewhat  marbled  with  yellow  and  moderate  body  crest  scales.  Compared 
yellow-green,  the  upper  surfaces  of  the  with  multistruppus,  caeruleolatus  has  mod- 
heads  were  pale  fawn,  the  eyeskin  pale  ally  4  (rather  than  2)  snout  scales  at  the 
grayish  green,  and  the  dewlaps  orange  in  level  of  the  second  canthals,  8  rather  than 
both  sexes.  7  vertical  rows  of  loreals,  5/5  rather  than 

The  series  of  A.  b.  caeruleolatus  includes  4/4   scales   between   the   interparietal   and 

four  juveniles   and  subadults   with   snout-  the  supraorbital   semicircles,   and   a   lower 

vent  lengths  from  60  to  91;  the  largest  of  mean  of  vertical  dorsal  scales  (17.1  versus 

these  is  a  topotype  that  was  colored  and  18.6).    With  regard  to  body  crest  scales, 


HisPANioLAN  Giant  Angle  •  Schwartz         129 


these  two  subspecies  show  the  same  situa- 
tion as  caeruleolatus  and  baleatus.  Com- 
pared with  .suJ)limis,  caeruleolatus  has  4 
(rather  than  2)  snout  scales  at  the  level  of 
the  second  canthals,  8  (rather  than  7)  ver- 
tical rows  of  loreals,  5/5  (rather  than  4/4) 
scales  between  the  interparietal  and  the 
supraorbital  semicircles,  and  lower  means 
of  both  vertical  dorsals  (17.1  versus  19.2) 
and  ventrals  (22.4  versus  25.1).  A.  h.  suh- 
limis  has  not  been  recorded  as  having  the 
dorsal  body  crest  scales  other  than  high, 
in  contrast  to  the  strong  tendency  in  cae- 
ruleolatus of  having  these  scales  moderate 
to  low. 

Discussion.  A.  h.  caeruleolatus  centers 
in  the  extremely  mesic  eastern  portion  of 
the  Valle  de  Cibao  in  that  area  that  has 
the  most  rainfall  in  the  Republica  Domin- 
icana.  I  have  already  commented  on  the 
specimens  from  Los  Bracitos,  Duarte  Prov- 
ince; these  specimens  are  old  and  pattern- 
less  and  are  from  a  locality  in  the  Cordillera 
Septentrional  which  is,  farther  west,  occu- 
pied by  A.  h.  baleatus;  I  include  them  with 
caeruleolatus  provisionally.  The  specimen 
from  Ponton,  Duarte  Province,  is  a  juve- 
nile (ASFS  V2987;  snout-vent  60)  and  is 
presently  patternless;  no  color  data  are 
available.  It  too  I  only  provisionally  re- 
gard as  caeruleolatus.  The  two  specimens 
from  La  Vega  Province  (ASFS  V4317, 
MCZ  128379)  are  also  without  color  data 
in  life,  and  the  former  is  a  patternless  ju- 
venile (snout-vent  69).  Specimens  from 
these  last  two  localities  also  require  verifi- 
cation as  to  subspecfic  status. 

A.  h.  caeruleolatus  presumably  inter- 
grades  with  four  subspecies:  baleatus,  mul- 
tistruppus,  the  subspecies  on  the  Peninsula 
de  Samana,  and  subspecies  to  the  south- 
east. Only  in  the  last  case  are  specimens 
that  I  interpret  as  intergradient  known, 
and  they  will  be  discussed  under  the  de- 
scription of  the  southeastern  subspecies. 
No  intergrades  are  known  between  the 
Samana  subspecies,  baleatus,  or  multistrup- 
pus.  Distance  between  caeruleolatus  and 
the  nearest  localities  for  these  subspecies 
are:     Samana     subspecies — 13     kilometers 


(Caiio  Abajo  and  5  km  NW  Sanchez); 
baleatus — 50  kilometers  (Los  Bracitos  and 
Pena);  multistruppus — 12  kilometers  (12.8 
km  NW  Bonao  and  Guaigiii).  Of  these 
presumed  areas  of  contact,  that  between 
caeruleolatus  and  the  Samana  subspecies  is 
not  unexpected;  the  area  between  the  two 
known  localities  is  very  open  and  relatively 
barren  and  devoid  of  trees  and  appears  al- 
ways to  have  been  so.  There  are  fine  high 
swamp-forests  in  the  western  part  of  this 
intervening  region,  and  it  is  possible  that 
intergrades  between  these  two  distinctive 
subspecies  will  be  encountered  in  these 
forests.  Most  puzzling  is  the  absence  of 
intergradation  between  caeruleolatus  and 
multistruppus.  The  specimen  from  north- 
west of  Bonao  is  a  juvenile,  but  it  does  not 
show  the  characteristic  multiple  crossbands 
of  both  young  and  adult  multistruppus.  It 
may  be  that  multistruppus  occupies  only 
the  foothills  of  the  Cordillera  Central  and 
that  the  zone  of  intergradation  between 
multistruppus  and  caeruleolatus  is  very 
abrupt. 

Remarks.  A.  b.  caeruleolatus  is  known 
from  sea  level  to  an  elevation  of  1000  feet 
(305  meters)  in  the  haitises  region  near 
Gonzalo  and  1200  feet  ( 366  meters )  north- 
west of  Bonao.  Specimens  were  secured 
primarily  from  native  collectors;  the  long 
series  of  topotypes  is  due  to  the  industry 
of  the  inhabitants  of  Cafio  Abajo.  The 
Cafio  Abajo  area  is  one  of  cafetales  and 
cacaotales  with  high  canopied  shade-trees, 
and  the  lizards  apparently  are  extremely 
abundant  in  this  optimal  habitat.  The 
pair  of  lizards  from  La  Mata  were  secured 
by  me  while  they  were  copulating  on  the 
side  of  a  large  shade-tree  in  a  cafetal  about 
4  feet  (1.2  meters)  above  the  ground  at 
1225  hours.  The  two  females  from  Gon- 
zalo were  taken  during  the  day  on  large 
trees  adjacent  to  a  small  spring  in  the 
haitises;  the  surrounding  area  was  under 
heavy  cultivation,  but  the  doline  slopes 
were  covered  locally  with  undisturbed  for- 
est. 

The  name  caeruleolatus  is  from  the 
Latin    "caeruleus"    for    "blue"    and    "latus" 


130         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


for  "side,"  in  allusion  to  the  sky-blue  lower 
sides  of  this  subspecies. 

Anolis  baleatus  samanae 
new  subspecies 

Holotype.  CM  54105,  an  adult  male, 
from  7.6  mi.  (12.2  km)  NE  Sanchez,  1000 
feet  (305  meters),  Samana  Province,  Re- 
publica  Dominicana,  one  of  a  series  col- 
lected by  native  collectors  on  28  November 
1971.   Original  number  ASFS  V34474  . 

Paratypes.  ASFS  V34475-79,  same  data 
as  holotype;  USNM  193990-92,  same  local- 
ity as  holotype,  native  collectors,  27  No- 
vember 1971;  MCZ  125634,  5.0  mi.  (8.0 
km)  NW  Sanchez  Province,  Republica  Do- 
minicana, J.  Aria,  27  November  1971;  ASFS 
V34495-96,  5.0  mi.  (8.0  km)  NW  Sanchez, 
Samana  Province,  Republica  Dominicana, 
J.  Aria,  28  November  1971;  CM  54127-30, 
5.0  mi.  (8.0  km)  NW  Sanchez,  Samana 
Province,  Republica  Dominicana,  J.  Aria, 
30  November  1971;  MCZ  12563.5-39, 
USNM  193993-4001,  5.0  mi.  (8.0  km)  NW 
Sanchez,  Samana  Province,  Republica  Do- 
minicana, J.  Aria,  1  December  1971;  ASFS 
V34514,  ASFS  V34836-38,  Las  Terrenas, 
Samana  Province,  Republica  Dominicana, 
native  collector,  28  November  1971;  ASFS 
V1904,  6  km  E  Sanchez,  Samana  Province, 
Republica  Dominicana,  R.  Thomas,  30  Oc- 
tober 1963;  AMNH  28651,  Samana,  Sa- 
mana Province,  Republica  Dominicana,  J. 
King,  August  1924;  AMNH  39817-23, 
AMNH  42285,  Laguna,  Samana  Province, 
Republica  Dominicana,  W.  G.  Hassler,  Oc- 
tober-December 1929;  USNM  61928,  Cayo 
Hondo,  Samana  Province,  Republica  Do- 
minicana, W.  L.  Abbott,  February  1919. 

Definition.  A  subspecies  of  A.  baleatus 
characterized  by  the  combination  of  mod- 
ally  2  snout  scales  at  level  of  second  can- 
thai  scales,  7  vertical  rows  of  loreal  scales, 
3  scales  between  the  interorbital  semicir- 
cles, 4/4  scales  between  the  inteiparietal 
and  the  supraorbital  semicircles,  moderate 
number  of  vertical  dorsal  scales  ( 13-20; 
mean  16.6),  moderate  number  of  ventral 
scales  (16-29;  mean  22.1),  nuchal  crest 
scales  very  high  to  high  (usually)  to  mod- 


erate or  low  (rarely)  in  both  sexes,  body 
crest  scales  high  to  moderate  but  often  low 
in  both  sexes,  subocular  scales  almost  al- 
ways separated  from  supralabial  scales  by 
one  (rarely  2)  row  of  scales;  dorsum  in 
both  sexes  in  life  blotched  dark  green, 
greenish,  dull  gray-green,  brown,  or  black- 
ish, dewlaps  in  males  dull  yellow  to  pale 
yellowish  orange,  in  females  very  pale  yel- 
low to  pale  yellowish  orange,  streaked  with 
blackish  or  brown  basally,  and  chin  and 
throat  in  males  cream  to  yellowish  or  yel- 
low-orange, mottled  with  black  or  gray,  in 
females  pale  green  to  greenish  yellow  with 
dark  green  to  brown  streaking  or  even  re- 
ticulate. 

Distribution.  The  Peninsula  de  Samana 
in  the  northeastern  Republica  Dominicana, 
and  apparently  islets  in  the  Bahia  de  Sa- 
mana. 

Description  of  holotype.  An  adult  male 
with  a  snout-vent  length  of  145  and  a  tail 
length  of  222  (regenerated);  snout  scales 
between  second  canthals  3,  6  vertical  rows 
of  loreal  scales,  3  scales  between  the  supra- 
orbital semicircles,  4/4  scales  between  the 
interparietal  and  the  supraorbital  semicir- 
cles, vertical  dorsals  18,  horizontal  dorsals 
19,  ventrals  21,  one  row  of  scales  between 
the  suboculars  and  supralabials,  fourth  toe 
lamellae  on  phalanges  II  and  III  30,  nuchal 
crest  scales  very  high,  body  crest  scales 
high;  in  life,  dorsum  mottled  dull  greens 
and  gray-brown  with  whitish  (almost 
cream  but  suffused  with  pale  gray);  upper 
surface  of  head  mixed  dark  brown  and 
gray,  venter  dull  greenish,  dewlap  orange, 
chin  and  throat  creamy  to  yellowish,  not 
marked  with  green. 

Variation.  The  series  of  54  A.  b.  sama- 
nae consists  of  32  males  and  22  females. 
The  largest  male  (AMNH  39807)  has  a 
snout-vent  length  of  157;  the  largest  fe- 
males (CM  54130,  USNM  193994)  have 
snout-vent  lengths  of  145.  The  male  is 
from  Laguna,  the  females  from  5.0  mi.  NW 
Sanchez.  Snout  scales  at  the  level  of  the 
second  canthals  range  between  2  and  5; 
the  mode  is  2  (24  specimens).  The  verti- 
cal loreal  rows  vary  between  5  and  9;  the 


HisPANioLAN  Giant  Angle  •  Scliwartz         131 


mode  is  7  (25  specimens).  There  are  2  or 
3  scales  between  the  supraorbital  semicir- 
cles (mode  3).  There  are  modally  4  scales 
between  the  interparietal  and  the  supraor- 
bital semicircles;  4  scales  are  involved  in 
43  percent  of  the  combinations:  actual 
counts  are  3/3  (2),  3/4  (3),  4/4  ( 17),  4/5 
(8),  5/5  (13),  5/6  (5),  6/6  (1),  6/7  (2), 
4/6  (1),  and  3/5  (1).  Vertical  dorsals 
range  between  13  and  20  (mean  16.6), 
horizontal  dorsals  between  13  and  27 
(19.3),  and  ventrals  16-29  (22.1).  Of  30 
adult  males,  14  have  the  nuchal  crest  scales 
very  high,  15  have  them  high,  and  one  has 
them  moderate;  in  20  adult  females,  nine 
have  the  nuchal  crest  scales  very  high, 
nine  have  them  high,  one  has  them  mod- 
erate, and  one  has  them  low.  Body  crest 
scales  in  males  are  high  in  three  lizards, 
moderate  in  16,  and  low  in  ten;  in  females, 
11  have  these  scales  moderate  and  ten  have 
them  low.  The  suboculars  are  separated 
from  the  supralabials  by  one  row  of  scales 
in  all  but  four  specimens  (7  percent), 
which  have  them  in  contact,  and  one  spec- 
imen (2  percent),  which  has  2  rows  of 
scales  in  this  position. 

A.  ]).  samanae  is  basically  a  blotched  liz- 
ard, and  no  adults  show  any  indication  of 
crossbanding.  The  body  is  irregularly 
blotched  with  blackish,  dark  green,  dull 
green,  gray-brown,  and  occasionally  there 
are  sky-blue  areas  along  the  ventrolateral 
margin  of  the  dorsal  coloration  in  males, 
but  these  areas  are  not  so  prominent  as  in 
caeruleolatus.  Regardless  of  the  dorsal 
shades,  the  upper  surface  of  the  head  is 
mixed  dark  brown  and  shades  of  gray  in 
both  sexes.  The  hindlimbs  are  finely 
barred  with  pale  and  dark  green.  The 
venter  is  dull  greenish  in  both  sexes.  The 
dewlap  in  males  varies  from  dull  yellow  or 
pale  yellowish  orange  to  orange,  and  the 
chin  and  throat  are  yellowish,  cream,  or 
yellow-orange,  mottled  with  black  or  gray. 
In  females,  the  dewlaps  are  very  pale  yel- 
low, pale  yellow-orange,  or  grayish  orange, 
at  times  streaked  with  blackish  or  brown 
basally,  and  the  chin  and  throat  ground 
color  is  pale  green,  marbled,  streaked,  or 


even  reticulate  with  dark  green  to  (rarely) 
brown. 

There  arc  one  juvenile  (AMNH  28651; 
snout-vent  length  40)  and  two  subadult 
(snout-vent  lengths  92  and  97)  A.  1).  sa- 
manae.  The  subadults  are  old  and  discolored 
but  their  patterns  seem  not  to  differ  from 
those  of  full  adults.  The  juvenile  on  the 
other  hand,  has  fom-  bold  pale  crossbars  on 
the  dorsum,  the  pattern  continuing  onto 
the  tail.  This  young  individual  has  the 
umbilicus  still  present  and  is  presumably 
near  hatchling  size. 

Comparisons.  Since  samanae  and  cae- 
ruleolatus are  adjacent  geographically,  the 
most  pertinent  comparisons  are  between 
them.  Examples  of  these  two  populations, 
as  noted  in  the  introduction  to  the  present 
paper,  were  available  to  me  simultaneously 
and  I  was  struck  with  their  differences  in 
life.  A.  1).  samanae  is  a  blotched  lizard 
whereas  caeruleolatus  is  a  crossbanded 
one;  the  latter  subspecies  also  typically  has 
sky-blue  ventrolateral  blotches,  a  feature 
absent  (or  occasionally  poorly  expressed) 
in  male  samanae.  Male  dewlap  colors  are 
similar  in  both  subspecies,  although  fe- 
male dewlap  colors  in  samanae  seem  some- 
what paler  than  those  of  caeruleolatus. 
The  chin  and  throat  markings  of  the  two 
subspecies  are  quite  distinct;  in  male  cae- 
ruleolatus, the  throat  is  deep  yellow  to 
yellow-orange,  at  best  with  very  faint  gray- 
ish dots  or  smudges,  whereas  in  male 
samanae  the  throat  is  yellowish  or  cream 
to  yellow-orange,  mottled  with  black  or 
gray.  In  female  caeruleolatus,  the  throat 
is  yellow  to  yellow-green,  always  with 
some  dark  green  dots,  blotching,  or  mar- 
bling, whereas  in  samanae  females,  the 
throat  is  pale  green,  greenish  yellow,  or 
yellow-green,  with  dark  green  to  brown 
streaking  or  reticulum. 

The  only  subspecies  thus  far  described 
which  is  blotched  like  samanae  is  the  Cor- 
dillera Central  suhlitnis,  although  caerul- 
eolatus may  show  a  marbled  dorsum  in 
some  areas.  No  pigmental  or  pattern  dif- 
ferences separate  samanae  and  sublimis, 
since  in  both  dorsal  coloration  and  color  of 


132         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


the  dewlap  the  major  color  involved  is 
green.  However,  the  throat  in  male  sub- 
limis  is  pale  green,  whereas  in  samanae  it 
is  cream  to  yellow-orange.  Certainly  rnulti- 
struppus  and  samanae  are  easily  distin- 
guished in  the  field  by  their  very  different 
dorsal  patterns,  for  example,  and  haleatus, 
with  its  very,  very  bright  chin  and  thioat, 
both  of  which  are  immaculate,  is  quite  dis- 
tinctive from  samanae. 

In  meristic  data,  samanae  differs  from 
caeruleolatus  in  having  2  (rather  than  4) 
snout  scales,  7  (rather  than  8)  vertical 
rows  of  loreals,  and  4/4  (rather  than  5/5) 
scales  between  the  inteiparietal  and  the 
supraorbital  semicircles.  From  multistrup- 
pus,  samanae  differs  in  having  a  lower 
mean  of  vertical  dorsal  scales  ( 16.6  versus 
1S.6),  and  the  same  difference  occurs  be- 
tween samanae  and  su])Umis  ( 16.6  versus 
19.2)  and  in  ventrals  (22.1  versus  25.1). 
From  haleatus,  samaiuie  differs  in  having 
2  (rather  than  4)  snout  scales,  4/4  (rather 
than  5/5)  scales  between  the  inteiparietal 
and  the  supraorbital  semicircles,  and  lower 
means  in  both  vertical  dorsals  ( 16.6  versus 
17.5)  and  ventrals  (22.1  versus  23.8).  The 
nuchal  crest  scales  in  samanae  are  more 
consistently  very  high  to  high  than  they 
are  in  any  of  the  other  subspecies  of  A. 
haleatus. 

Discussion.  As  pointed  out  in  the  dis- 
cussion of  A.  h.  caeruleolatus,  there  are  no 
intergrades  known  between  that  subspe- 
cies and  samanae.  The  isthmus  of  the  Pe- 
ninsula de  Samana  is  much  cleared  and 
locally  even  barren,  but  there  are  large 
western  swampy  areas  that  support  mag- 
nificent hardwood  forests  toward  the  land- 
ward side.  These  forests  may  well  support 
intermediates  between  samanae  and  cae- 
ruleolatus, or,  because  of  their  proximity 
to  the  mainland,  they  may  be  inhabited  by 
caeruleolatus.  Specimens  from  5.0  mi.  NW 
Sanchez,  that  locality  for  samanae  which 
is  closest  to  a  known  locality  for  caeruleo- 
latus ( 18  kilometers ) ,  show  no  tendencies 
toward  the  crossbanded  condition  of  cae- 
ruleolatus. 


A.  h.  samanae  is  the  only  Hispaniolan 
giant  anole  known  by  specimens  from  any 
off-shore  island  or  islet.  The  specimen 
from  Cayo  Hondo,  taken  by  W.  L.  Abbott, 
constitutes  this  record,  although  I  am  un- 
able to  locate  this  islet.  I  assume  it  is  one 
of  the  archipelago  within  the  Bahia  de  Sa- 
mana. 

Remarks.  All  but  one  A.  h.  samanae  se- 
cured by  myself  and  parties  were  native- 
collected.  The  exception  is  a  lizard  taken 
by  Richard  Thomas,  one  of  two  seen  on  a 
small  tree  and  in  a  vine  tangle  in  a  steep 
limestone  ravine  east  of  Sanchez.  The  area 
of  the  type  locality  is  in  the  uplands  of  the 
Sierra  de  Samana  on  the  road  between 
Sanchez  and  Las  Terrenas.  Thus  newly 
constructed  road  passes  through  superb 
mesic  high-canopied  forest,  and  much  of 
the  area  is  not  yet  seriously  disturbed.  Ob- 
viously from  the  number  of  lizards  secured 
by  natives  in  this  region,  A.  /;.  samanae  is 
common.  The  range  is  not  high,  with  a 
maximum  elevation  of  1673  feet  (510  me- 
ters) in  Monte  Las  Caiiitas;  this  mountain 
lies  between  Sanchez  and  Las  Terrenas. 
Specimens  from  Las  Terrenas  itself  were 
secured  by  natives  from  near-coastal  mesic 
cafetales  and  cacaotales,  and  lizards  from 
northwest  of  Sanchez  were  in  similar  situ- 
ations. 

Only  three  other  reptiles  (Diplo^lossus 
sternurus  alloeides  Schwartz,  Leiocephalus 
personatus  pyrrholaemus  Schwartz,  and 
Dromicus  parvifrons  niger  Dunn)  are  known 
to  have  differentiated  at  the  subspecific 
level  on  the  Peninsula  de  Samana.  Sphaero- 
clactylus  clenchi  Shreve  and  Sphaeroclac- 
tijlus  samanensis  Cochran  both  occur  there 
and  have  as  yet  unnamed  populations,  one 
of  which  in  each  case  is  limited  to  the  pen- 
insula. It  is  also  of  interest  to  note  that  in 
Anolis  clisticJius  Cope,  the  Samana  popula- 
tion is  identical  to  the  population  on  the 
soutliern  shores  of  the  Bahia  de  Samana 
(ignigularis  Mertens),  but  that  the  range  of 
this  subspecies  is  interrupted  at  the  head 
of  the  Bahia  de  Samana  by  A.  cl.  domini- 
censis  Reinhardt  and  Liitken  (see  Schwartz, 
1968:  280-81,  for  details). 


HisPANioLAN  Giant  Angle  •  Sclucmiz         133 


Anolis  baleatus  litorisilva  new  subspecies 
Holotype.  USNM  193977,  an  adult 
male,  from  1.2  km  SSW  Piinta  Cana,  La 
Altagracia  Province,  Rcpviblica  Domini- 
cana,  one  of  a  series  collected  by  Danny 
C.  Fowler  and  Bruce  R.  Sheplan,  on  24 
November  1971.  Original  number  ASFS 
V35095. 

Paratypes.  ASFS  \'35096-100,  same 
data  as  holotvpe;  CM  54113-14,  MCZ 
125616-17,  5.5  km  SSW  Punta  Cana,  La 
Altagracia  Province,  Republica  Domini- 
cana,  D.  C.  Fowler,  27  November  1971; 
ASFS  V29090,  Juanillo,  La  Altagracia 
Province,  Republica  Dominicana,  native 
collector,  24  July  1971;  ASFS  V961-62,  0.5 
mi.  NW  Boca  de  Yuma,  La  Altagracia 
Province,  Republica  Dominicana,  R.  F. 
Klinikowski,  R.  Thomas,  2  September  1963; 
ASFS  VI 136,  2.5  km  NW  Boca  de  Yuma, 
La  Altagracia  Province,  Republica  Domin- 
icana, native  collector,  4  September  1963; 
ASFS  V17573,  4  km  NW  Boca  de  Yuma, 
La  Altagracia  Province,  Republica  Domi- 
nicana, A.  Schwartz,  13  June  1969;  ASFS 
V17616,  2  km  NW  Boca  de  Yuma,  La  Al- 
tagracia Province,  Republica  Dominicana, 
J.  B.  Strong,  15  June  1969. 

Definition.  A  subspecies  of  A.  I)(ileatus 
characterized  by  the  combination  of  2  or  4 
scales  at  level  of  the  second  canthal  scales, 
7  vertical  rows  of  loreal  scales,  3  scales  be- 
tween the  interorbital  semicircles,  4/5 
scales  between  the  interparietal  and  the 
supraorbital  semicircles,  low  number  of 
vertical  dorsals  (13-19;  mean  15.9),  low 
number  of  ventral  scales  ( 18-26;  mean 
21.3),  nuchal  crest  scales  always  very  high 
to  high  in  both  sexes,  body  crest  scales 
high  (rarely)  to  moderate  or  low,  suboc- 
ular  scales  usually  separated  from  supra- 
labial  scales  by  one  row  of  scales;  dorsum 
in  life  varying  from  light  blue-brown  to 
light  greenish  brown  in  males,  dull  brown 
to  olive-brown  in  females,  blotched  with 
creamy  to  gray,  dewlap  in  males  bright 
orange,  brownish  in  females,  and  chin  and 
throat  (including  lips)  bright  orange  in 
males,  pale  yellow-green  in  females. 


Distri])ution.  Extreme  eastern  Repub- 
lica Dominicana  in  La  Altagracia  Province, 
from  Punta  Cana  to  the  \'icinity  of  Boca  de 
Yinna. 

Description  of  holotype.  An  adult  male 
with  a  snout-vent  length  of  136  and  a  tail 
length  of  183  (regenerated);  snout  scales 
between  sc^cond  canthals  2;  6  vertical  rows 
of  loreal  scales,  3  scales  between  the  supra- 
orbital semicircles,  5/5  scales  between  the 
interparietal  and  the  supraorbital  semicir- 
cles, vertical  dorsals  15,  horizontal  dorsals 
16,  ventrals  25,  subocular  scales  in  contact 
witli  the  supralabial  scales,  fourth  toe  la- 
mellae on  phalanges  II  and  III  31,  nuchal 
crest  scales  very  high,  dorsal  body  crest 
scales  high;  in  life,  dorsum  blotched  light 
blue-brown  and  light  green-brown;  venter 
pale  gray-green;  chin,  lips,  and  dewlap 
bright  orange. 

Variation.  The  series  of  16  A.  h.  litori- 
silva is  composed  of  six  males  and  ten  fe- 
males. The  largest  male  (MCZ  125616) 
has  a  snout-vent  length  of  158,  the  largest 
female  (ASFS  V961)  131.  The  male  is 
from  5.5  km  SSW  Punta  Cana,  the  female 
from  0.5  mi.  NW  Boca  de  Yuma.  Snout 
scales  at  the  level  of  the  second  canthals 
range  between  2  and  5;  there  are  t\vo 
modes,  2  and  4,  each  with  five  individuals. 
The  vertical  loreal  rows  vary  betwe(Mi  6 
and  9;  the  mode  is  7  (nine  specimens). 
There  are  2  to  4  scales  between  the  supra- 
orbital semicircles  (mode  3).  There  are 
modally  4/5  scales  between  the  interpari- 
etal and  the  supraorbital  semicircles;  5 
scales  are  involved  with  59  percent  of  the 
combinations;  actual  counts  are  4/4  (4), 
4/5  (6),  5/5  (5),  and  5/6  (1).  Vertical 
dorsals  range  between  13  and  19  (mean 
15.9),  horizcmtal  dorsals  between  14  and 
22  (18.5),  and  ventrals  between  18  and  26 
(21.3).  Of  four  adult  males,  three  have 
the  nuchal  crest  scales  very  high  and  one 
has  them  high;  of  five  adult  females,  two 
have  these  scales  very  high  and  three  have 
them  high.  In  the  males,  the  body  crest 
scales  are  high  in  one  and  moderate  in 
three,  and  in  the  females,  these  scales  are 


134         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


moderate  in  two  and  low  in  three.  The 
siiboculars  are  separated  from  the  siipra- 
labials  by  one  row  of  scales  in  all  but  one 
specimen  (6  percent). 

A.  h.  litorisilva  is  essentially  a  blotched 
lizard  whose  colors  do  not  include  bright 
or  even  medium  greens.  The  color  notes 
on  the  holotype  apply  equally  well  to  the 
other  adult  males — the  dorsum  is  blotched 
with  bluish  browns  and  light  greenish 
browns,  without  any  clear  greens,  and 
the  blotching  is  often  more  pronounced  on 
the  head  than  on  the  body.  In  females,  the 
dorsum  is  dull  brown  to  olive-brown  with 
only  occasional  slight  remnants  of  a  lighter 
green  pattern  on  the  head;  the  blotching 
in  the  female  involves  creamy  to  gray  pig- 
mentation. The  venter  is  pale  gray-green 
or  whitish  green  in  males,  pale  greenish 
gray  in  females.  The  dewlap  in  all  adult 
males  was  recorded  as  bright  orange,  and 
brownish  in  females.  In  males,  the  chin 
(including  the  lips)  is  bright  orange,  and 
pale  yellow-green  in  females.  The  upper 
surface  of  the  head  in  males  is  blotched 
like  the  body  and  is  dark  chocolate  in  fe- 
males. In  females,  the  upper  surfaces  of 
the  hindlimbs  were  recorded  as  olive- 
brown,  blotched  with  cream  to  gray  like 
the  dorsum. 

The  series  of  A.  h.  litorisilva  contains 
seven  juveniles  and  subadults  (snout-vent 
lengths  45  to  88).  The  smallest  juvenile 
(ASFS  V17573,  female)  was  bright  green 
in  life  with  four  pale  buffy  crossbands  and 
dark  green  shadow-bars  between  the  cross- 
bands;  the  tail  was  ringed  cream  and  dark 
gray,  and  the  venter  was  pale  green.  The 
dewlap  was  yellow-green  and  gray.  A 
slightly  larger  female  (ASFS  V17616)  with 
a  snout-vent  length  of  57  was  yellow-green 
dorsally  and  without  bands,  the  head  was 
brown;  the  eyeskin  was  green,  and  the 
venter  yellow-green.  The  tail  was  banded 
black  and  yellow-green,  and  the  dewlap 
was  mainly  brown  with  the  scale  rows  yel- 
low-green. A  still  larger  female  (ASFS 
V1136)  with  a  snout-vent  length  of  67  was 
green,  faintly  crossbarred  with  grayish 
green,  and  there  were  charcoal  smudges  on 


the  neck.  Two  male  subadults  with  snout- 
vent  lengths  of  71  and  83  (ASFS  V35099- 
100)  from  the  type  locality  were  recorded 
by  Fowler  as  follows:  "One  with  a  strong 
vertical  banding  pattern  alternating  brown- 
green  and  white-gray,  which  extends  from 
tip  of  tail  to  the  head  where  it  becomes 
slightly  more  diffuse;  on  the  other,  the  dor- 
sal groimd  color  is  dull  brown  with  rem- 
nants of  banding  pattern  only  around  head; 
the  ventral  ground  color  of  the  first  is  gray- 
green  with  brown  mottling,  the  second  is 
dull  gray-brown;  in  both  juveniles,  the 
dewlap  is  orange-green  and  the  chin  and 
lips  are  green."  The  largest  subadult 
(ASFS  V29090)  was  patternless  green 
above,  and  the  dewlap  was  orange  with 
charcoal  stripes;  the  specimen  is  a  female. 
Comparisons.  Because  of  its  blotched 
(rather  than  crossbanded)  pattern,  litori- 
silva requires  comparison  with  samanae 
and  suhlimis.  The  general  effect  of  the 
dorsa  of  all  three  subspecies  is  quite  sim- 
ilar, but  samanae  and  suhlimis  are  much 
the  brighter  lizards,  with  greens  predomi- 
nant in  the  dorsal  pigmentation.  On  the 
other  hand,  litorisilva  is  a  much  more  drab 
lizard,  without  clear  greens  in  the  adults, 
the  tendency  being  toward  more  sombre 
hues,  primarily  shades  of  browns.  From 
all  other  described  subspecies,  litorisilva 
differs  in  being  blotched  rather  than  cross- 
banded  and  also  in  having  much  less 
gaudy  dorsal  colors.  In  meristic  counts, 
litorisilva  differs  from  the  remaining  sub- 
species in  the  following  ways.  From  cae- 
ruleolattis,  litorisilva  differs  in  having  7 
(rather  than  8)  vertical  loreal  rows,  and 
lower  means  of  vertical  dorsals  (15.9  ver- 
sus 17.1)  and  ventrals  (21.3  versus  22.4). 
From  rnultistruppus,  litorisilva  differs  in 
lower  means  of  vertical  dorsals  ( 15.9  ver- 
sus 18.6)  and  ventrals  (21.3  and  22.3). 
From  stihlitiiis,  litorisilva  differs  in  having 
lower  means  of  vertical  dorsals  ( 15.9  ver- 
sus 19.2)  and  ventrals  (21.3  versus  25.1). 
From  haleatus,  litorisilva  differs  in  having 
lower  means  of  vertical  dorsals  ( 15.9  ver- 
sus 17.5)  and  ventrals  (21.3  versus  23.8). 
Meaningful  comparisons  of  litorisilva  with 


HisPANiOLAN  Giant  Anole  •  ScJiwartz         135 


other  subspecies  in  counts  of  snout  scales, 
and  scales  between  the  interparietal  and 
the  supraorbital  semicircles,  are  impossible 
since  litorisilva  has  a  bimodal  condition  in 
the  former  (and  the  bimodes  are  2  and  4, 
those  counts  which  occur  singly  as  the 
mode  in  the  other  subspecies)  and  has  a 
mode  of  4/5  in  the  latter  (whereas  all 
other  species  have  either  4/4  or  5/5). 
Considering  the  fairly  large  series  of  litori- 
silva (16  specimens),  these  two  "abnormal" 
conditions  are  puzzling.  At  least  in  the 
case  of  4/5  counts,  the  absence  of  3/3  or 
3/4  counts  in  litorisilva  suggests  that  this 
subspecies  tends  toward  a  5/5  count. 

Discussion.  A.  /;.  litorisilva  appears  to 
be  the  extreme  eastern  isolate  of  the  more 
widespread  A.  haleatus  stock.  It  occupies 
semi-arid  forests  on  and  near  the  coast  (as 
at  Juanillo  and  Punta  Cana)  and  on  the 
limestone  ridge  behind  Boca  de  Yuma. 
Both  situations  are  far  more  xeric  than  is 
customary  for  A.  haleatus,  and  the  faded 
nongreen  coloration  of  the  adults  is  doubt- 
less a  response  to  the  dry  and  open  to 
dense  forest  conditions  of  this  region. 
Nevertheless,  individuals  are  quite  con- 
spicuous at  night  as  they  sleep  exposed.  A. 
b.  litorisilva  presumably  comes  into  con- 
tact with  the  subspecies  to  the  north  and 
west  (named  below)  but  intergrades  are 
presently  unknown;  in  the  vicinity  of  Hig- 
iiey  (the  closest  locality  for  the  adjacent 
subspecies)  the  lizards  are  more  brightly 
colored  and  crossbanded  and  quite  unlike 
litorisilva. 

Remarks.  All  but  one  specimen  of  li- 
torisilva  were  collected  by  myself  and  par- 
ties. Individuals  were  found  sleeping  in 
primarily  coastal  forest  (to  which  the 
name,  from  "litus"  for  "shore"  and  "silva" 
for  "forest,"  refers  in  Latin)  at  elevations 
from  4  to  15  feet  (1.2  to  4.6  meters)  above 
the  ground.  Generally,  juveniles  sleep 
closer  to  the  groimd  and  in  more  dense 
situations  than  adults.  One  juvenile  was 
taken  from  a  roadside  Acacia,  a  most  un- 
usual situation  (since  Acacia  is  a  distinct 
xerophyte)  for  any  giant  anole.  Several 
adults   were  taken  in   dense  viny  tangles, 


sleeping  on  the  woody  vines;  the  advan- 
tage of  this  situation  was  made  (piite  ob- 
vious wIkmi  I  attempted  to  catch  a  large 
adult  at  night  by  hand.  The  light  from  my 
flashlight  wakened  the  lizard  almost  im- 
mediately, and  although  1  was  extremely 
careful  not  to  jar  any  of  the  vines,  this  was 
a  vain  endeavor.  At  the  first  jostling,  the 
lizard  jumped  to  the  ground  and  escaped 
in  the  dry  leaf  litter  and  understory. 

Anol'is  haleatus  scelestus  new  subspecies 

Holotype.  CM  54106,  an  adult  male, 
from  5.1  mi.  (8.2  km)  E  Santo  Domingo 
(from  Rio  Ozama),  Distrito  Nacional,  Re- 
publica  Dominicana,  one  of  three  collected 
by  David  C.  Leber  and  Richard  Thomas 
on  18  June  1964.  Original  number  ASFS 
V2460. 

Paratopes.  ASFS  V2461-62,  same  data  as 
holotype;  MCZ  125618-27,  8.4  mi.  (13.4 
km)  NE  La  Romana,  100  feet  (31  meters). 
La  Romana  Province,  Republica  Domini- 
cana, B.  R.  Sheplan,  22  November  1971; 
CM  54115-18,  USNM  193981-89,  8.4  mi. 
(13.4  km)  NE  La  Romana,  100  feet  (31 
meters).  La  Romana  Province,  Republica 
Dominicana,  D.  C.  Fowler,  A.  Schwartz, 
17  July  1971;  MCZ  16321,  La  Romana,  La 
Romana  Province,  Republica  Dominicana, 
E.  Leider,  1922;  ASFS  V29284-300,  0.2  mi. 
(0.3  km)  N  Otra  Banda,  350  feet  (107  me- 
ters). La  Altagracia  Province,  Republica 
Dominicana,  D.  C.  Fowler,  A.  Schwartz, 
26  July  1971;  ASFS  V21699-700,  1  km  NE 
Higiiey,  La  Altagracia  Province,  Republica 
Dominicana,  J.  R.  Dennis,  R.  Thomas,  16 
August  1969;  USNM  193979-80,  0.7  mi. 
(1.1  km)  W  Higiiey,  La  Altagracia  Prov- 
ince, Republica  Dominicana,  R.  Thomas, 
29  August  1963;  ASFS  V1038,  1  mi.  (1.6 
km)  W  Higiiey,  La  Altagracia  Province, 
Republica  Dominicana,  R.  Thomas,  3  Sep- 
tember 1963;  ASFS  V28757,  15.5  mi.  (24.8 
km)  E  San  Pedro  de  Macoris,  Rio  Cumay- 
asa.  La  Romana  Province,  D.  C.  Fowler, 
12  July  1971;  ASFS  V28910-16,  15.5  mi. 
(24.8  km)  E  San  Pedro  de  Macoris,  Rio 
Cumayasa,  San  Pedro  de  Macoris  Province, 
Republica   Dominicana,   D.   C.   Fowler,  A. 


136         Bulletin  Musewyi  of  Comparative  Zoology,  Vol.   146,  No.  2 


Schwartz,  16  July  1971;  ASFS  V28847,  15.5 
mi.  (24.8  km)  E  San  Pedro  de  Macoris,  La 
Romana  Province,  Repiiblica  Dominicana, 
A.  Schwartz,  15  July  1971. 

Associated  specimens.  REPUBLICA 
DOMINICANA:  La  Altagracia  Province,  1 
km  SE  Las  Lisas  (ASFS  V17434-35);  San 
Cristobal  Province,  8  km  N  Yamasa,  200 
feet  (61  meters)  (ASFS  V28656). 

Definition.  A  subspecies  of  A.  haleatus 
characterized  by  the  combination  of  mod- 
ally  2  scales  at  level  of  the  second  canthal 
scales,  7  vertical  rows  of  loreal  scales,  3 
scales  between  the  supraorbital  semicircles, 
5/5  scales  between  the  interparietal  and 
the  supraorbital  semicircles,  low  number  of 
vertical  dorsals  ( 12-20;  mean  15.4 ) ,  low 
number  of  ventral  scales  ( 17-28;  mean 
21.1),  nuchal  and  body  crest  scales  always 
very  high  to  high  in  both  sexes,  subocular 
scales  usually  separated  from  supralabial 
scales  by  one  (occasionally  two)  row  of 
scales;  dorsum  in  both  sexes  either  green 
with  three  pastel  green  crossbands  or  dark 
green  flecked  with  light  green,  cream  with 
some  greenish  to  brownish  green  smudges, 
dewlap  in  males  deep  yellow  to  deep  or- 
ange, streaked  or  smudged  with  dark 
brown  to  charcoal,  and  throat  in  females 
dark  green  marbled  with  yellow  and  pale 
green  (males  unrecorded). 

Distribution.  Southeastern  Republica 
Dominicana,  from  the  Sierra  de  Yamasa 
and  the  vicinity  of  Santo  Domingo  in  the 
west,  east  to  the  region  about  Higiiey  and 
Las  Lisas  in  La  Altagracia  Province. 

Description  of  Jiolotype.  An  adult  male 
with  a  snout-vent  length  of  152  and  a  tail 
length  of  267;  snout  scales  between  second 
canthals  4;  8  vertical  rows  of  loreal  scales, 
2  scales  between  the  supraorbital  semicir- 
cles, 4/5  scales  between  inteiparietal  and 
supraorbital  semicircles,  vertical  dorsals 
16,  horizontal  dorsals  16,  ventrals  22,  sub- 
ocular  scales  separated  from  supralabial 
scales  by  one  row  of  scales,  fourth  toe  la- 
mellae on  phalanges  II  and  III  34,  nuchal 
crest  scales  high,  body  crest  scales  moder- 
ate;   in   life,    dorsum   olive-green   with   six 


pastel  green  crossbands,  tail  and  venter 
light  green;  dewlap  dark  yellow. 

Variation.  The  series  of  61  A.  b.  sceles- 
ttis  consists  of  27  males  and  34  females;  a 
large  number  of  the  specimens  are  juve- 
niles and  subadults.  The  largest  male 
(ASFS  V29284)  has  a  snout-vent  length 
of  180,  the  largest  female  (ASFS  V29286) 
147;  both  are  from  near  Otra  Banda.  Snout 
scales  at  the  level  of  the  second  canthals 
range  between  2  and  4;  the  mode  is  2  (32 
specimens).  The  vertical  loreal  rows  vary 
between  5  and  8,  with  a  mode  of  7  (25 
specimens).  There  are  1  to  4  scales  be- 
tween the  supraorbital  semicircles  (mode 
3).  There  are  modally  5/5  scales  between 
the  interparietal  and  the  supraorbital  semi- 
circles; 5  scales  are  involved  in  49  percent 
of  the  combinations;  actual  counts  are  3/4 
(2),  4/4  (14),  4/5  (14),  5/5  (17),  5/6 
(11),  6/6  (1)  and  4/6  (1).  Vertical  dor- 
sals range  between  12  and  20  (mean  15.4), 
horizontal  dorsals  between  15  and  25 
(18.8),  and  ventrals  between  17  and  28 
(21.1).  Of  11  adult  males,  nine  have  the 
nuchal  crest  scales  very  high  and  two  have 
them  high.  Of  16  adult  females,  nine  have 
these  scales  very  high  and  seven  have  them 
high.  Body  crest  scales  in  males  are  high 
in  two  lizards,  moderate  in  eight,  and  low 
in  three;  in  females,  the  body  crest  scales 
are  high  in  two,  moderate  in  eight,  and  low 
in  six.  Fifty-three  specimens  have  the  sub- 
oculars  separated  from  the  supralabials  by 
one  row  of  scales,  whereas  in  four  lizards 
( 7  percent )  these  scales  are  in  contact,  and 
in  two  lizards  (3  percent)  they  are  sepa- 
rated by  tv/o  rows  of  scales. 

In  general,  both  sexes  of  A.  b.  scelestiis 
show  a  pattern  of  about  six  or  seven  fine 
crossbands  that  are  often  obscured  by  dor- 
sal blotching.  Colors  are  shades  of  greens, 
with  brighter  green  the  base  color  and  the 
blotching  tending  toward  darker  shades. 
The  crossbands  are  lighter  pastel  shades  of 
green,  and  in  some  lizards  the  dorsal 
ground  color  is  olivaceous.  Another  vari- 
ant, which  is  somewhat  more  prevalent  in 
females,   is   an  olive  green  to   dark   green 


HisPANioLAN  Giant  Angle  •  Schwaiiz 


137 


dorsum,  flocked  with  pale  green.  Two  fe- 
males from  near  Higiiey  showed  still  an- 
other style  of  body  pattern  and  color,  with 
the  dorsal  ground  color  cream  with  some 
dark  green  to  brownish  green  snuidges, 
and  the  neck  with  alternating  pale  blue 
and  charcoal  markings,  the  pale  blue  mark- 
ings persisting  onto  the  cheeks.  In  males 
the  upper  surface  of  the  head  is  brown, 
and  in  females  it  is  mixed  brown  and 
green,  with  the  snout  and  supraocular 
scales  deep  green  in  some  lizards.  In  fe- 
males, the  chin  and  throat  are  dark  green, 
marbled  with  yellow  and  pale  green.  The 
dewlap  is  rather  \'ariable;  in  males  it  has 
been  recorded  as  dark  yellow  or  deep  yel- 
low to  orange  or  dark  orange,  whereas  in 
females  the  dewlap  varies  from  yellow  to 
dark  orange  with  dark  brown,  olivaceous, 
or  charcoal  streaking,  marbling,  or  smudg- 
ing. Although  there  are  no  color  notes  in 
life,  in  the  preserved  lizards  the  eyeskin  is 
regularly  pale  gray,  and  I  presume  that  in 
life  the  eyeskin  is  set  off  from  the  rest  of 
the  head  color  in  some  pigmental  fashion. 
Many  specimens  of  both  sexes  have  the 
lower  sides  tigroid  with  "stripes"  extend- 
ing conspicuously  onto  the  lateral  sides  of 
the  abdomen. 

There  are  34  juvenile  and  subadult  A.  b. 
sceJestus,  with  snout-vent  lengths  between 
46  (USNM  193989)  and  94  (ASFS 
V21699-700).  Three  juveniles  (snout-vent 
lengths  46-61)  have  umbilici  still  present. 
This  entire  suite  of  young  lizards  shows  a 
remarkable  diversity  in  dorsal  pattern. 
Even  small  specimens  may  be  either  uni- 
color  green  (usually  with  a  vertical  nuchal 
white  crescent  and  a  white  subocular 
spot ) ,  green  with  three  or  four  yellow  body 
bands,  or  there  may  be  many  more  bands 
resulting  from  the  inteiposition  of  pale 
body  bands  between  the  primary  pale 
body  bands.  One  specimen  (ASFS  \'29296; 
snout-vent,  length  70,  male )  has  both  pale 
body  bands  and  interstitial  pale  blotching, 
whereas  another  lizard  (MCZ  125621; 
snout-vent  length  86,  female)  already 
shows  the  adult  pattern  of  several  fine  pale 


crossbands  on  a  green  ground.  The  largest 
subadults,  however,  (ASFS  V21699-700; 
snout-vent  lengths  94,  male  and  female) 
are  both  presently  unicolor  and  show  no 
indications  of  the  adult  body  banding. 
That  a  single  juvenile  may  demonstrate  a 
pattern  change  is  shown  by  the  following 
notes  on  ASFS  V28757,  a  female  with  a 
snout-vent  length  of  54:  "Alive,  emerald 
green  with  about  foiu'  pale  yellow  cross- 
bands  on  body;  dead — seven  narrow  brown 
body  bands  which  are  hollowed,  and  the 
dorsal  groimd  color  now  pale  yellow- 
green."  The  dewlap  in  young  males  is  or- 
ange, in  young  females  from  dull  yellow 
streaked  with  charcoal  to  charcoal. 

Comparisons.  In  color  and  pattern,  A. 
h.  scelestus  differs  from  all  other  subspe- 
cies. No  other  named  population  has  six 
or  seven  narrow  dorsal  crossbands;  even 
multistnippus  is  much  more  conspicuously 
banded  than  scelestus  and  lacks  any  sort 
of  dorsal  blotching.  A.  h.  scelestus  is 
known  to  intergrade  with  more  northern 
caeruleolatus  and  is  presumed  to  meet  li- 
torisilva.  In  each  case,  there  is  no  difficulty 
distinguishing  the  adjacent  forms  chromat- 
ically. A.  h.  caeruleolatus  typically  has  (in 
males)  sky-blue  blotches  along  the  junc- 
tion of  the  dorsal  and  ventral  colors,  and 
is  prominently  crossbanded  with  three  dor- 
sal crossbands.  A.  h.  litorisilva  is  a  blotched 
lizard,  the  dorsal  colors  much  more  drab 
than  those  of  scelestus,  tending  toward 
browns  and  brownish  greens.  Perhaps 
scelestus  most  closely  resembles  multi- 
stnippus, but,  although  both  are  banded, 
the  bands  in  multistruppus  are  much  finer 
and  much  more  numerous  than  the  six  or 
seven  pale  dorsal  crossbars  in  scelestus. 

A.  h.  scelestus,  with  modally  2  snout 
scales,  differs  from  caeruleolatus,  which 
has  4  snout  scales.  In  having  7  vertical  lor- 
eal  rows,  scelestus  differs  from  caeruleola- 
tus, which  has  8  rows.  In  having  5/5  scales 
between  the  interparietal  and  supraorbital 
semicircles,  scelestus  differs  from  samanae, 
multistruppus,  and  .mhlimis,  all  of  which 
have  4/4.    A.  h.  scelestus  has  the  lowest 


138         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


mean  of  vertical  dorsals  ( 15.4 )  of  all 
named  subspecies,  being  most  closely  ap- 
proached by  litorisilva  (15.9).  A.  b.  sceles- 
tus  males  are  larger  than  those  of  any  other 
subspecies  ( 180  in  scelestus,  158  in  litori- 
silva, which  is  second  largest)  and  in  fact 
this  subspecies  exceeds  all  other  Hispanio- 
lan  giant  anoles  in  size,  being  most  closely 
approached  by  male  A.  r.  ricordi,  which 
reach  a  snout-vent  length  of  160. 

Discussion.  I  am  uncertain  that  all 
specimens  included  in  scelestus  should  be 
so  associated.  This  is  especially  true  of  the 
specimen  from  near  Yamasa  (ASFS 
V28656);  this  is  a  juvenile  male  and  its 
taxonomic  status  remains  somewhat  in 
doubt,  since  it  is  young.  It  is  also  possible 
that  specimens  from  Santo  Domingo  like- 
wise are  not  identical  with  more  eastern 
lizards,  although  the  two  samples  agree 
fairly  well. 

A.  b.  scelestus  and  A.  b.  caeruleolatus  in- 
tergrade  in  the  region  of  El  Seibo  Prov- 
ince; I  have  examined  the  following  mate- 
rial from  El  Seibo  which  I  consider 
intergradient:  3.5  mi.  (5.6  km)  S  Sabana 
de  la  Mar  (ASFS  X7877);  2.1  mi.  (3.4  km) 
N  El  Valle  (ASFS  X7861-62);  3  km  N  El 
Valle  (ASFS  V3157-58);  10.5  km  N  Hato 
Mayor  (ASFS  V35329-30).  This  series 
consists  of  three  juveniles  and  four  young 
adults  (with  snout-vent  lengths  between 
112  and  127).  The  single  adult  male 
(ASFS  X7877)  was  tannish  gray  in  life 
with  darker  brown  blotches,  a  pale  green 
venter,  and  an  orange  dewlap.  Two  adult 
females  (ASFS  X7861-62)  were  pale  pea- 
green  with  vertical  gray  bars,  the  upper 
surface  of  the  head  grayish  tan,  venter 
green,  and  dewlap  grayish  orange.  The 
lower  jaw  and  throat  were  green  mottled 
with  darker  green.  In  general  this  series 
seems  closer  to  caeruleolatus  than  to  scel- 
estus, but  the  male  lacks  sky-blue  ventro- 
lateral markings.  On  the  other  hand,  the 
vertical  gray  bars,  recorded  for  the  female, 
resemble  the  pattern  of  scelestus  rather 
than  that  of  female  caeruleolatus.  It  seems 
likely  that  caeruleolatus  and  scelestus  in- 
tergrade  in  this  region. 


Remarks.  Almost  all  ASFS  scelestus 
were  secured  while  the  lizards  were  asleep 
at  night.  Typical  situations  are  lowland 
cacaotales  and  cafetales  with  their  high 
canopied  shade-trees,  along  lowland 
streams  (as  at  Otra  Banda  and  Yamasa), 
and  in  woods  associated  with  limestone 
cliffs  (east  of  Santo  Domingo).  The  long 
series  from  the  Rio  Cumayasa  is  from  the 
high  riverine  woods  along  that  stream;  re- 
markably, we  secured  only  juveniles  and 
subadults  at  this  locality,  despite  three 
nocturnal  visits.  One  juvenile  from  this  lo- 
cality was  taken  on  the  exposed  branch  of 
an  Acacia  tree  along  an  open  road.  Per- 
haps the  most  remarkable  place  whence  A. 
b.  scelestus  has  been  taken  is  the  locality 
northeast  of  La  Romana.  This  place  is  a 
deep  and  well-wooded  ravine  through 
which  flows  a  clear  stream;  however,  the 
ravine  is  completely  surrounded  on  all 
sides  by  cane  fields,  and  the  ravine  woods 
are  completely  isolated  at  the  ravine  rim 
from  other  such  ecologies,  if  they  even  still 
exist  in  this  area.  A.  b.  scelestus  was  ex- 
ceptionally abundant  in  this  particular  and 
very  restricted  strip  of  riverine  gallery  for- 
est. Elevations  above  ground  recorded  for 
sleeping  scelestus  range  from  2  to  20  feet 
(0.6  to  6.1  meters),  with  juveniles  usually 
sleeping  much  lower  than  adults.  The  alti- 
tudinal  distribution  of  A.  b.  scelestus  is  in 
general  low,  with  recorded  elevations  from 
sea  level  to  200  feet.  It  is  likely  that  this 
subspecies  also  occurs  in  the  uplands  of 
the  Cordillera  Oriental,  but  as  yet  there 
are  no  specimens  from  areas  within  that 
rather  low-lying  but  mesic  and  well-for- 
ested massif. 

The  name  scelestus  is  from  the  Latin  for 
"unlucky,  wretched,"  in  allusion  to  the  dif- 
ficulties involved  with  collecting  this  sub- 
species at  the  La  Romana  ravine  noted 
above. 

The  transition  betu'cen  scelestus  and  //- 
torisilva  must  be  very  abrupt;  the  two  sub- 
species are  known  from  localities  separated 
by  only  28  kilometers.  The  habitats  of  the 
two  subspecies  are  quite  different,  with 
scelestus  inhabiting   very   mesic  situations 


HisPANiOLAN  Giant  Angle  •  Schwartz         139 


and  litorisilva  xeiic  coastal  woods.  Inter- 
estingly, this  same  eastern  region  of  the 
Repiiblica  Dominicana  is  also  an  area  of 
abrupt  changes  in  subspecies  of  Anolis  di- 
stichus,  where  the  subspecies  ifi,ni<iularis 
Mertens  and  properus  Schwartz  have 
ranges  which  coincide  rather  closely  with 
those  of  scelestus  and  litorisilva  (see 
Schwartz,  1968:  275,  map).  The  question 
of  intcrgradation  between  scelestus  and  the 
southwestern  subspcx-ies  next  to  be  named 
will  be  discussed  under  that  taxon. 

Perhaps  more  so  than  any  other  subspe- 
cies, scelestus  seems  to  show  a  very  spotty 
distribution.  Two  instances  are  worthy  of 
mention.  There  are  excellent  extensive 
coastal  forests  at  Cabo  Caucedo  south  of 
the  Aeropuerto  Internacional  de  las  Ame- 
ricas on  the  southern  Dominican  coast.  Re- 
peated diurnal  and  nocturnal  visits  to  these 
splendid  woods  yielded  no  A.  baleatus,  de- 
spite what  seems  to  be  more  than  adequate 
habitat.  A  second  locality,  east  of  Boca 
Chica  along  the  same  coast,  likewise  sup- 
ports extensive  fine  stands  of  lowland  hard- 
wood forests,  and  there  also,  despite  many 
diurnal  and  nocturnal  visits,  we  have  never 
encountered  A.  baleatus.  It  is  possible  that 
these  two  instances  of  fairly  dry  coastal 
woods  are  not  suitable  for  scelestus 
(whereas  they  surely  would  be  for  litori- 
silva) and  that  scelestus  simply  does  not 
occur  there. 

Anolis  baleatus  fraudator 
new  subspecies 

Holotype.  USNM  193978,  an  adult  fe- 
male, from  4  km  W,  6  km  N  Azua,  Azua 
Province,  Repiiblica  Dominicana,  one  of 
two  taken  by  Richard  Thomas,  on  23  July 
1969.   Original  number  V21384. 

Paratypes.  ASFS  V21385,  same  data  as 
holotype;  ASFS  V21433,  Barreras,  Azua 
Province,  Repiiblica  Dominicana,  native 
collector,  25  July  1969;  ASFS  V723,  1.1  mi. 
(1.8  km)  S  San  Jose  de  Ocoa,  1400  feet 
(427  meters),  Peravia  Province,  Repiiblica 
Dominicana,  R.  F.  Klinikowski,  24  August 
1958;  ASFS  V21203,  Sierra  Martin  Garcia, 
about  3000  feet  (915  meters),  above  Bar- 


reras, Azua  Province,  Repiiblica  Domini- 
cana, R.  Thomas,  20  July  1969;  ASFS 
V31207,  Sierra  Martin  Garcia,  above  Bar- 
reras, between  2000  and  2800  feet  (610 
and  854  meters),  west  slope,  Mt.  Biisi'i, 
Barahona  Province,  Repiiblica  Dominicana, 
B.  R.  Sheplan,  15-17  September  1971.' 

Definition.  A  subspecies  of  A.  baleatus 
characterized  by  the  combination  of  mod- 
ally  4  scales  at  level  of  the  second  canthal 
scales,  6  vertical  rows  of  loreal  scales,  2  or 
3  scales  between  the  supraorbital  semicir- 
cles, 4/4  scales  between  the  interparietal 
and  the  supraorbital  semicircles,  high  num- 
ber of  vertical  dorsal  scales  (17-21;  mean 
18.8),  low  number  of  ventral  scales  (18- 
26;  mean  20.7),  nuchal  scales  high,  body 
crest  scales  moderate  in  only  adult  female, 
subocular  scales  usually  separated  from 
supralabial  scales  by  one  row  of  scales; 
dorsum  (in  female)  mottled  pale  and 
darker  gray,  with  three  irregular  white 
crossbands,  and  blotched  with  yellow- 
green,  top  of  snout  and  lores  straw,  labials 
dull  yellow,  and  dewlap  nearly  white  with 
a  yellowish  or  cream  wash. 

Distribution.  The  Sierra  Martin  Garcia 
in  Barahona  and  Azua  provinces,  and 
along  the  southern  slopes  of  the  Cordillera 
Central  and  the  Sierra  de  Ocoa  in  Azua 
and  Peravia  provinces. 

Description  of  holotype.  An  adult  fe- 
male with  a  snout-vent  length  of  133  and 
tail  length  of  244;  snout  scales  between 
second  canthals  4;  6  vertical  rows  of  loreal 
scales,  2  scales  between  supraorbital  semi- 


1  Since  the  present  manuscript  was  completed, 
a  juvenile  female  ( MCZ  132301)  with  a  snout- 
vent  length  of  57  mm,  was  secured  by  E.  E.  Wil- 
liams and  J.  Roughgarden  at  a  locality  south  of 
La  Honna,  Peravia  Province,  on  19  July  1972. 
This  lizard  is  to  be  considered  a  paratype.  It  has 
3  snout  scales  at  the  level  of  the  second  canthals, 
6  loreal  rows,  3  scales  between  the  supraorbital 
semicircles,  5/5  scales  between  the  interparietal 
and  the  semicircles,  16  vertical  rows  of  dorsal 
scales  and  20  rows  of  ventral  scales,  and  1  scale 
between  the  suboculars  and  the  supralabials.  Both 
nuchal  and  body  crest  scales  are  low.  As  pre- 
served, the  lizard  is  dull  greenish  with  indications 
of  dark  dorsal  crossbars,  and  it  lacks  any  pale 
dorsal  markings. 


140         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


circles,  4/3  scales  between  the  interpari- 
etal and  the  supraorbital  semicircles,  verti- 
cal dorsals  17,  horizontal  dorsals  24, 
ventrals  21,  subocular  scales  separated 
from  supralabial  scales  by  one  row  of 
scales,  fourth  toe  lamellae  on  phalanges  II 
and  III  33,  nuchal  crest  scales  high,  body 
crest  scales  moderate;  in  life,  dorsum  mot- 
tled pale  and  dark  gray,  blotched  with  yel- 
low-green and  with  three  irregular  white 
crossbands,  labials  dull  yellow,  top  of  snout 
and  lores  straw,  and  dewlap  nearly  white 
with  a  yellowish  or  cream  wash. 

Variation.  The  only  adult  is  the  holo- 
type;  the  remainder  of  the  paratypic  series 
is  composed  of  juveniles  and  subadults 
with  snout-vent  lengths  between  74  and  96 
(three  males,  two  females).  Snout  scales 
at  the  level  of  the  second  canthals  range 
between  2  and  4;  the  mode  is  4  ( four  spec- 
imens). The  vertical  loreal  rows  vary  be- 
tween 5  and  7,  with  a  mode  of  6  (three 
specimens).  There  are  2  or  3  scales  be- 
tween the  supraorbital  semicircles;  both 
categories  have  the  same  frequency.  There 
are  modally  4/4  scales  between  the  inter- 
parietal and  the  supraorbital  semicircles;  4 
scales  are  involved  in  67  percent  of  the 
combinations;  actual  counts  are  3/4  (2), 
4/4  (3),  and  5/6  (1).  Vertical  dorsals 
range  between  17  and  21  (mean  18.8), 
horizontal  dorsals  between  20  and  24 
(21.4),  and  ventrals  between  18  and  26 
(20.7).  The  only  adult  specimen  (a  fe- 
male) has  the  nuchal  crest  scales  high  and 
the  dorsal  body  crest  scales  moderate.  Five 
specimens  have  the  suboculars  separated 
from  the  supralabials  by  one  row  of  scales 
and  one  lizard  has  these  scales  in  contact 
( 17  percent ) . 

The  details  of  the  color  and  pattern  of 
the  only  adult,  the  female  holotype,  have 
already  been  given.  The  juveniles  and 
subadults  show  the  same  general  pattern 
configuration  as  does  the  adult.  The  small- 
est juvenile  (snout-vent  length  72),  a  fe- 
male topotype,  was  gray  with  yellowish 
mottling  and  a  pat*"ern  of  three  irregular 
crossbands,  a  faint  white  scapular  stripe, 
and    black    postauricular    and    postorbital 


spots.  The  dewlap  was  charcoal  with 
white  scales.  The  next  largest  individual 
(snout-vent  length  74),  a  male,  had  the 
dorsum  pale  green  with  irregular  trans- 
verse barring;  the  upper  surfaces  of  the 
limbs  were  pale  green  and  gray-green,  and 
the  tail  was  banded  pale  green  and  gray- 
green.  The  venter  was  whitish.  The  dew- 
lap was  very  dark  yellowish  with  an  or- 
ange wash  posteriorly.  A  slightly  larger 
male  (snout-vent  length  85)  was  pale 
green,  much  marbled  and  shaded  with  tan 
to  gray  and  with  some  faint  evidence  of 
transverse  crossbands;  the  chin  and  throat 
were  gray-green,  and  the  dewlap  pale  gray- 
ish orange.  A  female  from  the  Sierra  Mar- 
tin Garcia  (snout-vent  length  88)  was 
green  and  brown  dorsally  and  without  pale 
markings;  the  dewlap  was  marbled  with 
charcoal.  Finally,  the  largest  subadult 
(snout-vent  length  76),  a  male,  had  the 
dewlap  dirty  yellow  with  orange  streaking. 
In  the  case  of  fraudator,  the  very  pale  ( al- 
most white)  adult  fem.ale  dewlap  appears 
to  be  preceded  ontogenetically  by  brighter 
and  more  typically  A.  haleatus  hues. 

Comparisons.  No  other  subspecies  of  A. 
haleatus  approaches  the  pale  colors  of 
fraudator,  nor  does  any  other  subspecies 
have  such  a  pale  dewlap.  Although  frau- 
dator combines  the  blotching  and  trans- 
verse crossbands  in  the  same  fashion  as 
does  scelestus,  fraudator  is  in  all  ways  a 
paler  lizard.  Comparisons  in  details  of 
color  and  pattern  with  all  other  subspecies 
of  A.  haleatus  are  unnecessary.  A.  h.  frau- 
dator differs  from  samaime,  scelestus,  multi- 
struppus,  and  suhliinis  in  having  4  rather 
than  2  snout  scales  at  the  second  canthals, 
and  only  fraudator  has  a  mode  of  6  verti- 
cal rows  of  loreals  (7  or  8  in  all  other  sub- 
species). In  having  4/4  scales  between  the 
inteiparietal  and  the  supraorbital  semicir- 
cles, fraudator  differs  from  caeruleolatus, 
scelestus,  and  haleatus,  all  of  which  have 
5/5.  Although  fraudator  has  a  high  mean 
(18.8)  of  vertical  dorsals,  in  which  it  is 
exceeded  only  by  suhlimis  (mean  19.2), 
fraudator  has  the  lowest  mean   (20.7)   of 


HisPANiOLAN  Giant  Angle  •   Schwartz 


141 


ventrals  of  all  subspecies,  being  approached 
most  closely  by  scelestus  (21.1). 

Discussion.  Apparently  A.  h.  fraudator 
is  a  pale  subspecies  that  is  restricted  to  fa- 
vored situations  in  the  xeric  regions  asso- 
ciated with  the  Llanos  de  Azua  along  the 
southern  slopes  of  the  Cordillera  Central 
and  the  Sierra  de  Ocoa,  a  southern  affiliate 
of  the  former  range.  The  subspecies  ap- 
parently also  occurs  in  the  Sierra  Martin 
Garcia,  an  eastern  isolate  of  the  Sierra  de 
Neiba  (which,  it  will  be  recalled,  is  else- 
where occupied  by  A.  r.  ricordi)  and  sur- 
rounded by  extreme  desert.  The  specimen 
from  Barreras,  which  lies  at  the  foot  of  the 
Sierra  Martin  Garcia,  is  interesting  in  that 
it  seems  a  most  unlikely  locality  for  any 
giant  anole;  however,  I  assume  that  the 
specimen,  which  was  native-collected,  was 
taken  either  in  nearby  Cocos  groves  or  on 
the  lower  wooded  slopes  of  the  range  it- 
self. Two  specimens  from  the  higher  eleva- 
tions of  the  Martin  Garcia  are  from  dense 
woods,  and  the  specimen  from  San  Jose 
de  Ocoa  was  taken  from  a  large  tree  at  the 
edge  of  a  pasture.  The  type  locality  is 
semi-xeric  woods  with  vine  tangles  and 
mango  trees  in  an  otherwise  cultivated  but 
xeric  region.  Probably  A.  b.  fraudutor  is 
widely  distributed  in  suitable  situations 
through  much  of  this  region,  but  the  lizard 
appears  to  be  rare;  Buffett  and  I  collected 
in  semi-mesic  riverine  woods  at  a  locality 
4  km  W  and  17  km  N  Azua  at  an  elevation 
of  about  500  feet  ( 153  meters ) ,  both  dur- 
ing the  day  and  at  night,  without  seeing 
any  giant  anoles.  Natives  just  south  of  San 
Jose  de  Ocoa  at  an  elevation  of  1400  feet 
(427  meters)  did  not  secure  specimens  for 
us  in  semi-mesic  woodlands.  Since  the  al- 
titudinal  distribution  of  fraudator  extends 
from  sea  level  to  about  3000  feet  (915  me- 
ters) in  the  Sierra  Martin  Garcia,  the  ele- 
vations of  the  above-mentioned  localities 
are  within  the  known  altitudinal  range  of 
the  subspecies,  and  indeed  our  San  Jose  de 
Ocoa  locality  was  quite  close  to  where 
Klinikowski  secured  one  of  the  paratypes. 

Remarks.  The  name  fraudator  is  from 
the  Latin  for  "deceiver"  in  reference  to  the 


resemblances  between  this  sui)species  and 
A.  harahonae.  In  fact,  my  decision  to  re- 
gard fraudator  as  a  subspecies  of  baleatus 
rather  than  harahonae  is  based  more  upon 
the  juveniles  than  the  adults  of  fraudator; 
this  is  not  exclusively  due  to  the  fact  that 
there  are  more  juveniles  of  fraudator  than 
adults  but  rather  that  the  patterns  shown 
by  juvenile  fraudator  are  more  typically 
those  of  A.  Jjaleatus  than  of  A.  harahonae. 
A.  J),  harahomie  and  A.  h.  fraudator  are 
alike  in  modal  numbers  of  scales  at  the 
level  of  the  second  canthals  (4),  vertical 
loreal  rows  (6),  and  scales  between  the  in- 
terparietal and  the  supraorbital  semicircles 
(4/4),  and  they  do  not  differ  strikingly  in 
means  of  body  scales  ( 17.2,  18.8  in  vertical 
dorsals;  18.2,  21.4  in  horizontal  dorsals; 
22.1,  20.7  in  ventrals).  In  these  means, 
harahonae  is  lower  in  dorsal  body  counts, 
but  higher  in  ventral  coimts.  The  moder- 
ate nuchal  crest  scales  of  fraudator  occur 
also  in  harahonae,  but  most  female  hara- 
honae have  these  scales  low.  No  female 
harahonae  has  moderate  dorsal  body  crest 
scales  as  does  the  female  fraudator, 
whereas  moderate  body  crest  scales  occur 
in  females  of  most  subspecies  of  A.  halea- 
ttis  ( only  female  suhhniis  lack  them ) .  Tak- 
ing all  evidence  into  consideration,  I  have 
elected  to  consider  fraudator  a  subspecies 
of  A.  ])aleatus,  but  its  resemblances  to  A. 
harahonae  are  acknowledged.  The  distance 
separating  these  two  species  in  this  area 
is  only  20  kilometers  (see  introduction), 
and  it  is  not  unlikely  that  A.  harahomie  has 
been  derived  from  fraudator  across  the 
strait  that  is  now  the  Valle  de  Neiba  (see 
discussion).  On  the  other  hand,  A.  h. 
fraudator  is  removed  by  some  60  kilometers 
from  the  nearest  A.  ricordi  locality  in  the 
nearby  Sierra  de  Neiba.  There  is  no  ques- 
tion that  fraudator  is  not  correctly  associ- 
ated nomenclatorially  with  A.  ricordi. 

The  apparent  geographic  isolation  of 
fraudator  in  relation  to  other  subspecies  of 
A.  haleatus  is  probably  artificial.  The  near- 
est records  for  other  subspecies  are:  cae- 
ruleolatus — 38  kilometers  ( San  Jose  de 
Ocoa  and  La  Cumbre);  scelestus — 55  kilo- 


142         Bulletin  Museum  of  Comparaiive  Zoology,  Vol.   146,  No.  2 


meters  (San  Jose  de  Ocoa  and  Yamasa); 
and  sublimis — 50  kilometers  (San  Jose  de 
Ocoa  and  south  of  EI  Rio).  There  are 
suitable  habitats  for  giant  anoles  between 
caertileolatus,  scelestus,  and  fraudator,  but 
specimens  are  lacking.  The  intervening 
high  Cordillera  Central  between  the  ranges 
of  sublimis  and  fraudator  probably  acts  as 
a  barrier  to  prevent  contact  between  these 
two  subspecies. 

DISCUSSION 

My  decision  to  consider  Anolis  ricordi 
as  three  species  rather  than  one  has  some 
precedent  in  the  Schwartz  and  Garrido 
(1972)  treatment  of  the  Cuban  Anolis 
equestris,  wherein  that  species  was  divided 
into  five  species.  However,  the  two  situa- 
tions, although  comparable,  are  far  from 
identical.  In  the  A.  equestris  complex, 
there  are  at  least  a  few  incidences  of  sym- 
patry  between  members  of  the  species- 
complex  which  give  clues  to  the  facts  of 
the  situation;  there  are  sti'ong  differences 
in  size  of  dorsal  scales;  there  are  some 
strong  differences  between  details  of  pat- 
tern and  coloration  of  the  axillary  stripe 
and  the  dewlap  which  likewise  suggest 
that  we  are  there  dealing  with  more  than 
one  species.  But  on  the  other  hand,  the 
Hispaniolan  giant  anoles  show  absolute 
differences  in  the  nuchal  and  body  crest 
scales  and  differences  in  the  pattern  of  the 
body  itself,  as  well  as  modal  differences  in 
scutellar  details.  In  addition,  there  are  no 
cases  as  yet  known  in  Hispaniola  of  sym- 
patry  between  the  three  entities  that  I  re- 
gard as  full  species.  The  gaps  between 
them  are  narrow,  however,  and  I  feel 
strongly  that  it  is  merely  a  matter  of  get- 
ting into  the  intermediate  areas  and,  once 
there,  being  fortunate  enough  to  encounter 
giant  anoles. 

It  should  be  obvious  from  my  systematic 
treatment  that  I  am  convinced  that  we  are 
dealing  in  Hispaniola  with  three  distinct 
species — ricordi,  barahonae,  and  baleatus. 
Surely  the  differences  between  ricordi  and 
baleatus  are  such  that,  when  taken  in  sum, 
one  has  no  doubts  that  he  is  involved  with 


two  very  different  animals.  The  differences 
here  are  much  greater,  for  instance,  than 
between  Anolis  distichus  Cope  and  Anolis 
hrevirostris  Bocourt,  two  species  that  were 
long  confused  and  that  resemble  each 
other  moiphologically  to  a  very  great  de- 
gree. Yet  once  one  learns  what  the  char- 
acters are  for  separating  them,  he  experi- 
ences little  difficulty  in  dealing  with  both 
populations  or  individuals,  either  alio-  or 
sympatric,  of  these  two  species.  The  dif- 
ferences in  life,  as  far  as  pattern  and  color 
are  concerned,  are  not  particularly  subtle, 
and  the  details  of  scutellation  are  not  di- 
chotomous,  but  the  modal  differences  are 
so  well  correlated  with  the  pigmental  and 
pattern  traits  that  we  now  recognize  these 
two  species  with  assurance. 

An  even  more  obvious  parallel  is  Anolis 
carolinensis  Voigt  and  Anolis  allisoni  Bar- 
bour in  Cuba.  These  two  species  of  green 
anoles,  long  confused  as  A.  porcatus  Gray 
(or  A.  c.  porcatus),  were  shown  by  Ruibal 
and  Williams  (1961)  to  be  a  sibling  pair, 
fairly  allopatric  but  both  widely  distrib- 
uted throughout  much  of  Cuba,  and  to 
differ  structurally  by  the  condition  of  the 
postauricular  area.  The  presence  (allisoni) 
or  absence  (carolinensis)  of  a  deep  and 
elongate  postauricular  groove  in  these  two 
species  is  correlated  with  very  striking  dif- 
ferences in  adult  pattern  and  coloration 
and  other  details  of  scutellation. 

The  same  situation,  that  of  two  species 
masquerading  under  a  single  name,  can 
also  be  demonstrated  in  Anolis  alutaceus 
Cope  and  Anolis  clivicola  Barbour  and 
Shreve  (Schwartz  and  Garrido,  1971),  and 
the  two  species  recently  confused  under 
Anolis  spectrum  Peters;  both  these  situa- 
tions pertain  to  Cuban  species.  Sr.  Gar- 
rido also  advises  me  that  he  has  much 
evidence  to  indicate  that  Anolis  cyanopleu- 
rus  cupeyalensis  Peters  is  in  fact  a  sym- 
patric sibling,  rather  than  a  subspecies,  of 
A.  cyanopleurus  Cope. 

I  could  cite  other  examples  in  Antillean 
iguanids  (Leiocephalus)  and  anguids 
(Diploglossus)  which  demonstrate  quite 
clearly  the  above  trend.    As  more  material 


HisPAXioLAN  Giant  Angle  •  Schwartz 


143 


from  more  diverse  localities  becomes  avail- 
able, and  as  this  material  is  subjected  to 
re-evaluation  with  differing  and  more  mod- 
ern philosophies,  our  impressions  of  rela- 
tionships among  Antillean  anolines  have 
been  modified  or  changed.  A  major  factor 
in  such  revisions  has  invariably  been  a 
great  quantity  of  new  material  from  areas 
that  had  previously  been  unsampled,  cou- 
pled with  pigmental,  ecological,  and  etho- 
logical  data  from  the  living  specimens.  A 
second  general  line  of  evidence,  equal  to 
or  possibly  suipassing  morphological  and 
distributional  data  in  importance,  is  karyo- 
typic  and  electrophoretic  information.  One 
or  both  of  these  areas  of  investigation  are 
increasing  our  knowledge  of  the  complex- 
ities within  such  a  genus  as  Anolis.  When 
these  two  areas  of  research — morphologi- 
cal and  biochemical — can  be  brought  to 
bear  simultaneously  upon  a  single  species 
or  species  complex,  the  results  may  be  even 
more  meaningful  than  either  is  alone.  As 
yet  this  has  not  been  done  in  any  of  the 
Antillean  giant  anoles,  so  that  my  conclu- 
sions, based  upon  morphology  and  distri- 
bution, remain  to  be  verified  by  other  evi- 
dence. Yet  I  feel  as  secure  as  any 
systematist  can  be  when  he  is  dealing  with 
data  that  are  incomplete. 

As  pointed  out  in  the  introduction  to  the 
present  paper,  the  taxa  ricordi,  haleatus, 
harahonae,  and  leberi  are,  on  inspection, 
unequivocally  distinct.  But  the  degree  or 
level  of  differentiation  of  these  four  taxa 
seems  to  be  two-fold.  On  one  hand  (ri- 
cordi and  haleatus),  the  two  populations 
are  easily  separable  on  the  basis  of  a  struc- 
tural feature  (the  nuchal  crest  scales),  a 
character  that  is  strongly  correlated  with 
obvious  pigmental  and  pattern  traits.  On 
the  other  hand,  the  differences  between 
harahonae  or  leheri  and  ricordi  are  pri- 
marily ones  of  pigmentation  and  pattern, 
with  morphological  differences  much  less 
trenchant  than  between  ricordi  and  ha- 
leatus. At  the  outset  such  a  dichotomy  sug- 
gests that  it  might  be  more  proper  to  con- 
sider "A.  ricordi"  as  a  complex  of  full 
species  than  as  one  species  with  four   (or 


more)  subspecies.  Appar(Mitly  Williams 
and  Rand  (1969)  had  the  same  inclina- 
tions, since  they  indicated  that  the  differ- 
ences between  some  of  the  then-named 
populations  of  A.  ricordi  were  such  as  to 
suggest  that  there  might  be  more  than  one 
species  involved. 

Once  the  above  assumption  has  been 
made — namely,  that  A.  ricordi  is  composed 
of  more  than  one  species — then  the  prob- 
lem first  becomes  one  of  differentiating  and 
delimiting  the  component  species.  There 
is  no  difficulty  here  in  separating  A.  ricordi 
and  A.  haleatus  on  the  basis  of  crest  scales. 
None  of  the  populations  of  A.  haleatus  has 
the  moderate  (rarely)  to  low  (usually) 
nuchal  crest  scales  of  A.  r.  ricordi.  In  ad- 
dition, the  narrow  geographical  gaps  that 
exist  between  A.  ricordi  and  A.  haleatus 
also  suggest  that  these  two  taxa  may  be 
either  allopatric  or  may  meet  and  occur 
sympatrically  without  intergradation. 

The  status  of  the  Tiburon  populations 
that  I  associate  nomenclatorially  with  A. 
ricordi  and  that  of  A.  harahonae  as  a  dis- 
tinct species  are  less  clear  than  the  ricordi- 
haleatus  relationship.  First,  the  named 
populations  leheri,  viculus,  and  suhsolamis 
have  in  common  a  suite  of  pattern  and 
color  features  that  ally  them  more  closely 
to  each  other  than  to  A.  r.  ricordi.  The 
only  evidence  for  this  relationship  is  the 
occurrence  of  presumed  viculus  X  ricordi 
intergrades  in  the  Miragoane-Paillant  re- 
gion. Were  it  not  for  these  specimens,  I 
would  be  strongly  tempted  to  consider  the 
three  Tiburon  taxa  as  comprising  a  sepa- 
rate species.  Any  interpretation  of  the  re- 
lationships of  the  Tiburon  taxa  suffers 
from  paucity  of  material  from  a  variety  of 
localities. 

The  situation  with  A.  harahomie  is  in 
some  ways  puzzling.  Although  there  is  no 
question  that  it  is  distinct  from  A.  ricordi, 
its  relationships  to  A.  haleatus  are  much 
less  certain.  This  uncertainty  is  caused  by 
A.  h.  fraudator,  that  population  assigned  to 
A.  haleatus  which  is  closest  geographically 
to  A.  harahonae.  It  is  particularly  unfortu- 
nate  that  fraudator   is   known   from   only 


144         Bulletin  Museum  of  Coinparative  Zoologij,  Yo].   146,  No.  2 


one  adult  and  several  juvenile  and  sub- 
adult  specimens,  since  adult  males  ( primar- 
ily) would  be  most  instructive  in  compar- 
ing frattdator  with  barahonae.  On  the 
other  hand,  the  closeness  of  fraiidator  and 
barahonae  in  characteristics  may  be  rather 
a  reflection  of  the  ancestry  of  A.  bara- 
honae— namely,  that  it  is  a  south  island 
(sensu  Williams,  1961)  invader  from  the 
north,  and  that  the  parent  population  has 
been  fraudator  rather  than  any  other  sub- 
species of  A.  baleatus  or  A.  ricordi  from  the 
west. 

It  might  be  more  proper  either  to  con- 
sider A.  barahonae  as  conspecific  with  A. 
baleatus  (the  two  taxa  linked  through 
fraudator) ,  or  to  consider  fraudator  a  sub- 
species of  A.  barahonae;  either  interpreta- 
tion has  merit.  The  course  that  I  have 
taken  seems  satisfactory  at  the  moment  but 
surely  is  subject  to  reinterpretation  with 
the  acquisition  of  more  material  from  this 
critical  geographic  area. 

The  history  of  the  Hispaniolan  giant 
anoles  appears  to  be  correlated  with  the 
two  palaeo-islands  that  have  been  fused  at 
the  level  of  the  Cul  de  Sac-Valle  de  Neiba 
plain  with  lowering  Pleistocene  sea  levels. 
I  suggest  the  following  history  for  the  com- 
plex; the  reader  should  keep  in  mind  that 
such  a  history  is  based  upon  taxonomic 
premises  that  are  inductive,  and  the  cau- 
tions and  uncertainties  that  I  expressed 
above  have  special  application  here. 

Distributional  evidence  suggests  that  the 
giant  Hispaniolan  anole  stock  was  origi- 
nally restricted  to  the  north  island  (north 
of  the  Cul  de  Sac-Valle  de  Neiba  plain). 
In  this  region,  two  distinctive  species  arose, 
ricordi  in  the  west  and  baleatus  in  the  east. 
There  apparently  has  been  local  differen- 
tiation on  the  north  island  at  a  subspecific 
level  in  both  these  species,  but  that  in  ri- 
cordi remains  unanalyzed  because  of  too 
few  specimens.  On  the  other  hand,  differ- 
entiation in  A.  baleatus  is  now  fairly  well 
known  and  documented.  This  species  oc- 
curs east  of  the  Cordillera  Central  and  on 
the  southern  slopes  of  that  range  and  in 
the  Sierra  Martin  Garcia.   There  have  been 


local  population  differentiations  in  response 
to  the  various  ecologies  within  the  area  oc- 
cupied, with  two  major  integumental 
trends  (coloration  and  pattern)  and  details 
of  scutellation  of  the  head  and  body  (al- 
though the  latter  is  not  so  clear  as  the  for- 
mer ) . 

There  seem  to  have  been  two  subse- 
quent invasions  of  the  south  island.  To  the 
west,  a  (presumably)  early  invasion  of  the 
A.  ricordi  stock  crossed  what  is  now  the 
Cul  de  Sac  Plain  into  the  Port-au-Prince 
area.  It  is  pertinent  that  many  north  island 
species  have  made  this  same  crossing  and 
have  extended  their  ranges  but  little  fur- 
ther. These  species  with  more  restricted 
ranges  have  been  handicapped  either  by 
competition  with  already  established  spe- 
cies, improper  ecological  situations,  or  rela- 
tively recent  arrival.  A.  ricordi  seems  to 
have  been  an  early  arrival,  without  local 
competitors,  and  with  abundant  proper 
ecology  (mesic  forests).  The  species  has 
thus  expanded  its  range  after  the  original 
crossing  to  cover  the  entire  Tiburon  Penin- 
sula, having  somewhere  succeeded  in  cross- 
ing the  mountainous  spine  of  the  Massif  de 
la  Hotte-Massif  de  la  Selle.  Local  differ- 
entiation along  the  Tiburon  in  response  to 
lack  of  genetic  contact  across  the  interior 
mountains  has  also  taken  place.  Further 
speculations  on  details  of  the  history  of  A. 
ricordi  on  the  Tiburon  Peninsula  are  point- 
less, since  the  specimens  upon  which  any 
generalisations  may  be  made  do  not  as  yet 
exist  in  collections. 

A  second  invasion  to  the  east  occurred 
presumably  at  a  later  date,  after  the  estab- 
lishment of  A.  ricordi  on  the  Tiburon  Pen- 
insula. This  latter  invasion  resulted  in  the 
differentiation  of  A.  barahonae  (from  a 
fraudator  or  pre-fraudator  stock  on  the 
southern  portion  of  the  north  island )  in  the 
Sierra  de  Baoruco  and  its  subsequent  ex- 
pansion onto  the  southern  portion  of  the 
Peninsula  de  Barahona  and  east  along  the 
southern  slopes  of  the  Sierra  de  Baoruco. 
With  the  previous  establishment  of  A.  ri- 
cordi to  the  west  (as  at  Thiotte),  the 
western   movement  of  A.    barahonae   was 


HisPANiOLAN  Giant  Angle  •  Schwartz 


145 


halted  by  the  presence  of  the  related  spe- 
cies. I  have  no  doubt  that  both  A.  ricordi 
and  A.  barahonae  will  be  found  to  be 
closely  allopatric  or  synipatric  in  extreme 
southeastern  Haiti  between  the  Dominico- 
Haitian  border  and  Saltrou,  and  also  that 
these  two  species  meet  and  interact  along 
the  northern  slopes  of  the  Sierra  de  Bao- 
ruco  and  the  Morne  des  Enfants  Perdus. 

One  other  distributional  detail  requires 
comment.  The  occurrence  of  A.  r.  ricordi 
in  the  main  mass  of  the  Sierra  de  Neiba  on 
the  northern  side  of  the  Valle  de  Neiba 
and  of  A.  h.  fraudator  in  the  Sierra  Martin 
Garcia,  an  extreme  eastern  isolate  of  the 
Sierra  de  Neiba,  has  already  been  noted. 
The  Martin  Garcia  seems  to  have  been 
long  isolated  from  not  only  the  Sierra  de 
Neiba  but  also  from  all  other  Hispaniolan 
mountain  masses;  it  is  ideally  a  montane 
island  in  a  sea  of  desert.  It  seems  likely 
that  this  range  was  unoccupied  by  giant 
anoles  of  either  species  (A.  ricordi  or  A. 
baleatus),  despite  the  fact  that  the  range 
forms  a  portion  of  the  Neiba  uplift.  Inva- 
sion of  the  Martin  Garcia  was  possible 
from  either  the  northwest  (ricordi)  or  the 
northeast  (baleatus).  Of  the  two  species, 
A.  baleatus  was  the  more  vagile  and 
reached  the  Sierra  Martin  Garcia  from  the 
relatively  more  mesic  southern  slopes  of 
the  Cordillera  Central  before  A.  ricordi 
reached  it  across  the  deserts  and  xeric  hills 
between  the  Sierra  de  Neiba  and  the  Mar- 
tin Garcia.  This  upland  population  in 
turn  was  responsible  for  the  invasion  of  the 
Sierra  de  Baoruco  across  the  better  for- 
ested and  more  mesic  eastern  end  of  the 
Valle  de  Neiba. 

Wetmore  and  Swales  (1931:  235)  re- 
ported the  finding  of  recent  Anolis  ricordi 
skeletal  material  in  Barn  Owl  (Tyto  alba) 
pellets  from  L'Acul,  Dept.  du  Sud,  Haiti, 
on  the  Tiburon  Peninsula,  and  Hecht 
(1951:  23)  noted  the  abimdant  remains  of 
the  species  from  deposits  in  "Deep  Cave," 
near  St.  Michel  de  I'Atalaye,  Dept.  de  I'Ar- 
tibonite,  Haiti.  Etheridge  (1965:  101)  re- 
ported A.  ricordi  remains  from  recent  owl 
pellets  near  the  mouth  of  a  cave  near  Boca 


de  Yuma,  La  Altagracia  Province,  Repu- 
blica  Dominicana.  Etheridge  (op.  cit.:  87- 
88)  also  noted  pre-Columbian  giant  anole 
remains  from  a  cave  at  Cerro  de  San  Fran- 
cisco near  Pedro  Santana,  La  Estrelleta 
Province,  Repiiblica  Dominicana.  From  the 
suite  of  about  80  cranial  elements  and 
eight  pelves,  Etheridge  extrapolated  that 
the  maximally  sized  individuals  in  the  cave 
deposits  had  a  snout-vent  length  of  190- 
192  mm,  some  30  mm  larger  than  any  liv- 
ing A.  ricordi  recorded  ( 159  mm,  fide  Eth- 
eridge, op.  cit.:  88).  The  maximally  sized 
Hispaniolan  giant  anole  recorded  in  the 
present  paper  reaches  a  length  of  180  mm 
(male  A.  b.  scelestus  from  Otra  Banda,  La 
Altagracia  Province,  Republica  Domini- 
cana). The  difference  between  this  mod- 
ern living  lizard  and  the  maximally  sized 
pre-Columbian  lizards  is  not  so  great  as 
Etheridge's  data  suggest.  Intriguingly,  the 
Cerro  de  San  Francisco  area  lies  within  the 
known  range  of  A.  r.  ricordi,  and  the  larg- 
est specimens  of  this  subspecies  ( male  with 
a  snout-vent  length  of  160  mm,  female  151 
mm)  are  from  the  southern  slopes  of  the 
Cordillera  Central,  veiy  close  to  the  Cerro 
de  San  Francisco  area.  Although  there 
seems  to  have  been  some  change  in  maxi- 
mum size  in  Hispaniolan  giant  anoles  with 
the  passage  of  time,  these  changes  have 
not  been  of  the  magnitude  that  previous 
data  suggested. 

LITERATURE  CITED 

Cochran,  D.  M.  1941.  The  herpetology  of 
Hispaniola.  Bull.  U.S.  Natl.  Mus.,  177:  398, 
120  figs.,  12  pis. 

Cope,  E.  D.  1864.  Contributions  to  the  herpe- 
tology of  tropical  America.  Proc.  Acad.  Nat. 
Sci.    Philadelphia,  pp.  166-181. 

DUMERIL,      A.      M.      C,     AND      G.      BiBRO.N.       1837. 

Erpetologie  gcnerale  ou  histoire  naturelle 
complete  des  reptiles,  vol.  4.  571  pp.,  14  pis. 

Etheridge,  R.  E.  1965.  Fossil  lizards  from  the 
Dominican  Republic.  Quart.  Jour.  Florida 
Acad.  Sci.,  28(1):  83-195,  3  figs. 

Garrido,  O.  H.,  AND  A.  Schwartz.  1968.  Cu- 
ban lizards  of  the  genus  Chamaeleolis.  Quart. 
Jour.  Florida  Acad.  Sci.,  30(3):  197-220,  2 
figs. 

Hecht,  M.  K.     1951.     Fossil  lizards  of  the  West 


146         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  2 


Indian  genus  Aristelliger  ( Gekkonidae ) . 
Amer.  Mus.  Novitates,  No.  1538:  1-33,  8  figs. 

Maerz,  a.,  and  M.  R.  Paul.  1950.  A  Diction- 
ary of  Color.  New  York:  McGraw-Hill  Book 
Co.,  pp.  vii  +  1-23,  137-108,  56  pis. 

Mertens,  R.  1939.  Herpetologische  Ergebnisse 
einer  Reise  nach  der  Insel  Hispaniola,  West- 
indien.  Abh.  senckenberg.  naturf.  Ges.,  449: 
1-84,  10  pis. 

Rand,  A.  S.,  and  E.  E.  Williams.  1969.  The 
anoles  of  La  Palma;  aspects  of  their  ecolog- 
ical relationships.  Breviora,  Mus.  Comp. 
Zool.,  No.  327:  1-18,  1  fig. 

RuiBAL,  R.,  AND  E.  E.  Williams.  1961.  Two 
sympatric  Cuban  anoles  of  the  carolinensis 
group.  Bull.  Mus.  Comp.  Zool.,  125(7): 
183-208,  11  figs. 

Schmidt,  K.  P.  1921.  Notes  on  the  herpetology 
of  Santo  Domingo.  Bull.  Amer.  Mus.  Nat. 
Hist.,  44(11):  7-20,  12  figs. 

Schwartz,  A.  1964.  Anolis  equestris  in  Ori- 
ente  Province,  Cuba.  Bull.  Mus.  Comp. 
Zool.,  131(12):  407-428,  7  figs. 

.     1968.     Geographic    variation    in    Anolis 

distichus  Cope  ( Lacertilia,  Iguanidae )  in  the 
Bahama  Islands  and  Hispaniola.  Bull.  Mus. 
Comp.  Zool.,  137(2):  255-309,  4  figs.,  2  pis. 

,  and  O.  H.  Garrido.     1971.     The  status 

of    Anolis    alutaceus    clivicolus    Barbour    and 


Shreve.    Caribbean  Jour.  Sci.,   11(1-2):    11- 
15. 

.     1972.     The  lizards  of  the 


AND 


Anolis  equestris  complex  in  Cuba.  Stud. 
Fauna  Curasao  and  Caribbean  Is.,  39(134): 
1-86,  8  figs. 

Thomas,  R.  1971.  A  new  species  of  Diploglos- 
sus  (Sauria:  Anguidae)  from  Hispaniola. 
Occ.  Papers  Mus.  Zool.,  Louisiana  State  Univ., 
40:   1-9,  4  figs. 

Wetmore,  A.,  and  B.  H.  Swales.  1931.  The 
birds  of  Haiti  and  the  Dominican  Republic. 
Bull.  U.S.  Natl.  Mus.,  155:  1-483,  2  figs.,  26 
pis. 

Williams,  E.  E.  1961.  The  evolution  and  rela- 
tionships of  the  Anolis  semilineatus  group. 
Breviora,  Mus.  Comp.  Zool.,  No.  138:  1-8,  1 
pi. 

.  1962.  Notes  on  Hispaniolan  herpetol- 
ogy. 6.  The  giant  anoles.  Breviora,  Mus. 
Comp.  Zool.,  No.  155:  1-15,  1  fig. 

.  1965.  Hispaniolan  giant  anoles  (Sau- 
ria, Iguanidae ) :  new  data  and  a  new  subspe- 
cies. Breviora,  Mus.  Comp.  Zool.,  No.  232: 
1-7,  2  figs. 

,  and  a.  S.  Rand.  1969.  Anolis  insolitus,  a 

new  dwarf  anole  of  zoogeographic  importance 
from  the  mountains  of  the  Dominican  Repub- 
lic. Breviora,  Mus.  Comp.  Zool.,  No.  326: 
1-21,  6  figs. 


us  ISSN  0027-4100 


BulLetln  OF  TH 


seum 


Comparative 
Zoology 


A  Revision  of  the  Cardinalfish  Genus 

Epigonus  (Perciformes,  Apogonidae)^ 

with  Descriptions  of  Two  New  Species 


GARRY  F.  MAYER 


HARVARD  UNIVERSITY 

CAMBRIDGE,  MASSACHUSETTS,  U.S.A. 


VOLUME  146,  NUMBER  3 
19  SEPTEMBER  1974 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED   BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 


Breviora.  1952- 
BULLETIN   1863- 

Memoirs  1864-1938 

JoHNsoNiA,  Department  of  Mollusks,  1941- 

OccAsioNAL  Papers  on  Mollusks,  1945- 

SPECIAL  PUBLICATIONS. 

1.  Whittington,  H.  B.,  and  E.  D.  I.  Rolfe  (eds.),   1963.    Phylogeny  and 
Evolution  of  Crustacea.  192  pp. 

2.  Turner,  R.  D.,  1966.  A  Survey  and  Illustrated  Catalogue  of  the  Teredini- 
dae  (Mollusca:  Bivalvia).  265  pp. 

3.  Sprinkle,  J.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echinoderms. 
284  pp. 

4.  Eaton,  R.  J.  E.,  1974.  A  Flora  of  Concord.  211  pp. 

Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.    Fishes  of  the  Gulf  of  Maine. 
Reprint. 

Brues,  C.  T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.    Classification  of 
Insects. 

Creighton,  W.  S.,  1950.   The  Ants  of  North  America.    Reprint. 

Lyman,   C.   P.,   and  A.   R.   Dawe    (eds.),    1960.    Symposium   on   Natural 
Mammalian  Hibernation. 

Peters'  Check-list  of  Birds  of  the  World,  vols.  2-7,  9,  10,  12-15. 

Proceedings  of  the  New  England  Zoological  Club  1899-1948.    (Complete 
sets  only.) 

Publications  of  the  Boston  Society  of  Natural  History. 

Price  list  and  catalog  of  MCZ  publications  may  be  obtained  from  Publications 
Office,  Museum  of  Comparative  Zoology,  Harvard  University,  Cambridge,  Massa- 
chusetts, 02138,  U.S.A. 

©  The  President  and  Fellows  of  Harvard  College  1974. 


t 


A  REVISION  OF  THE  CARDINALFISH  GENUS  EPIGONUS 
(PERCIFORMES,   APOGONIDAE),   WITH    DESCRIPTIONS 
OF  TWO  NEW  SPECIES^ 


GARRY  F.   MAYERS 


TABLE    OF    CONTENTS  resim-ected  and   redescribed  on  the  basis   of  new 

.                                                                                    ,^Y  material,   and  Htjnnodus  atherinoides  Gilbert   and 

:^        j*^     " ,  ,«  H.  megalops  Smith  and  Radcliffe  are  synonymized 

Introduction 14^  ^^^^^j^  ^   occidentaUs  Goode  and  Bean. 

Metnods     Species   descriptions   include   discussions   of  dis- 

Systematics    """r " ici  tribution,       geographic       variation,       ontogenetic 

Genus  Epigonus  Rafmesque  151  ^^^^^^^    a„j    taxonomic    problems.     An    in%'estiga- 

Diagnosis   151  ^^^^  ^^  ^j^^  ^^p^^  ^^j  ^    lenimen  (Whitley)  reveals 

Description    151  ^^^^^  ^^  holotype  and  paratypes   are  not   conspe- 

Key  to  the  Species  of  Epigonus 152  ^.jjj^     Instead,  the  paratypes  are   members  of  E. 

Epigonus  telescopus  152  denticulatus  Dieuzeide.    A  key  to  the  species   of 

Epigomis   macrops  159  Epigonus    is    provided    at    the    beginning    of   the 

E))igonus  ))andionis    163  paper. 

Epigonus  fragilis  169 

Epigonus  occidentaUs 170  INTRODUCTION 

Epigonus  denticulatus   1^5 

Epigonus  oligolepis 179  Selected  species  of  Epigomis  have  been 

Epigonus  tretvavasae  183  j^^^^^^^^^  1^    ^^^.^^^  f^^  at  least  two  hundred 

Epigonus  pectimfcr    loo  J                     , ,  onr,     r>^\                i     j 

Epigonus  wbustus  1S9  ti^ty  Y^ars.    Vaillant   (1888:  25)   remarked 

Epigonus  lenimen    193  that  E.  telescopus  was  recognized  in  ancient 

Epigonus  crassicaudus  197  times,  and  Risso  (1810:  303)  reported  that 

Species  Incertae  Hedis 199  j-j-jj^  gpecies  was  prized  for  its  firm,   deli- 

Aek„S^!Sr°!!!' ::;::::::::::::::::::::::::::  loE!  cious-tasti„g  „,«,,  although  it  was  rardy 

Literature  Cited 200  taken.   The  presence  of  common  names  tor 

Appendix 203  E.    telescopus   in   vocabularies    of   western 

Mediterranean  and  North  Atlantic  fishing 

Abstr.\ct.  a  study  of  the  deep-sea  Apogonidae  communities  ( Doderlein,  1889 )  provides 
results  in  a  revision  of  the  genus  Epigonus  additional  evidence  of  man's  long-term 
Rafinesque.  Twelve  species  are  recognized  in-  awareness  of  the  species.  £.  telescopus  is 
eluding  two  new  forms — E.  oligolepis  and  £...,,  •  n  i  i  •  .  i  i  i.  „r 
pectirUfer.  E.  fragilis  (Jordan  and  Jordan)  is  still  occasionally  sold  in  the  markets  of 
southwestern  Europe. 

'  This  paper  is  based  on  a  portion  of  a  thesis  Two  other  species  of  EpigOnus  are  cap- 
presented  to  Harvard  University  in  partial  ful-  tured  by  commercial  fishermen.  E.  denti- 
fillment    of    the    requirements    for    the    Ph.D.    in  culcitUS   is    edible    ( Dieuzeide    et   al.,    1953: 

^i?'"g>-                ^                                                  r  -1^)    and  is   taken  in  the   Mediterranean. 

-  Department   of   Marine   Science,    University   of  y-r    ..,               .i      ,i  .      ,■                          ^^t■^^■  >.,    f^^ 

c    u  \ri     J      c^    u  .     I          171    •  1     oo-m        1  Uutil  recently  this  form  was  mistaken  tor 

South  Florida,   St.   Petersburg,   Florida  33/01   and  ^■'^_,        ,                     _,              ,          , 

Museum    of   Comparative   Zoology,    Harvard    Uni-  the  yOUng  of  E.  teUsCOpUS.    E.  crassicaudus 

\ersity,  Cambridge,  Massachusetts  02138.  is    caught   by   Chilean   fishermen.     Like   E. 

Bull.  Mus.  Comp.  Zool.,  146(3):  147-203,  September,  1974        147 


148       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


telescopus,  it  is  not  taken  in  sufficient 
numbers  to  support  a  separate  fishery  but  is 
captured  by  fishermen  trawhng  for  more 
abundant  deepwater  organisms. 

Although  African  Epigonus  are  not  pres- 
ently exploited,  tropical  eastern  Atlantic 
stocks  may  represent  future  sources  of  pro- 
tein for  mankind.  Surveys  sponsored  by 
the  Organisation  of  African  Unity  and  the 
U.S.  Agency  for  International  Development 
revealed  these  fishes  are  "of  possible  po- 
tential importance  ( not  necessarily  by  pres- 
ent marketing  standards)  [Williams,  1968: 
79]."  The  same  may  be  true  for  Caribbean 
and  Gulf  of  Mexico  Epigonus;  however, 
complete  data  have  not  been  compiled  for 
the  latter  areas. 

A  major  hindrance  to  the  evaluation  of 
deep-sea  cardinalfish  stocks  has  been  taxo- 
nomic  confusion.  The  systematic  history  of 
Epigonus  began  in  1810  with  Risso's  de- 
scription of  Pomatomus  telescopus  and 
Rafinesque's  account  of  its  synonym  Epi- 
gonus niacrophthahnus.  During  the  fol- 
lowing seventy-one  years,  work  on  the 
genus  was  primarily  limited  to  re-descrip- 
tions of  E.  telescopus  and  discussions  of  its 
biology  (e.g.,  Cuvier,  1828;  Valenciennes, 
1830;  Capello,  1868;  Moreau,  1881). 

The  surge  in  oceanographic  exploration 
during  the  last  quarter  of  the  nineteenth 
and  beginning  of  the  twentieth  centuries 
rapidly  increased  the  number  of  nominal 
Epigonus AAke  species.  Among  the  forms 
described  between  1881  and  1920  were 
Apogon  pandionis  Goode  and  Bean,  1881; 
E.  occidentalis  Goode  and  Bean,  1896; 
Hynnodus  atherinoides  Gilbert,  1905;  Oxij- 
odon  macrops  Brauer,  1906;  and  Hynnodus 
me  galops  Smith  and  Radcliffe,  1912.  In 
the  following  decade,  three  new  species 
and  two  new  genera  appeared  in  the  liter- 
ature. 

Much  of  the  confusion  associated  with 
the  taxonomy  of  Epigonus  stems  from  ma- 
terial described  prior  to  1930.  Early  taxa 
were  based  on  small  samples.  Because 
many  nations  participated  in  oceanographic 
research,     specimens     were     deposited    in 


scattered  institutions  and  descriptions  ap- 
peared in  diverse  publications.  Conse- 
quently, it  was  difficult  for  workers  to 
obtain  either  comparative  material  or  a 
broad  overview  of  the  group's  systematics. 
These  shortcomings  were  aggravated  by 
inaccurate,  under-illustrated  descriptions 
based  on  ill-considered  characters.  It  was 
common,  for  example,  to  use  dentition 
patterns  to  define  generic  boundaries,  yet 
tooth  arrangements  are  difficult  to  observe, 
easily  damaged,  and  subject  to  ontogenetic 
and  geographic  variation.  As  a  result,  an 
inordinately  large  number  of  Epigonu.s-\ike 
forms  was  recognized  by  the  end  of  the 
1920's. 

Although  generic  taxonomy  was  stream- 
lined by  Fowler  and  Bean  in  1930  and 
Matsubara  in  1936,  species-level  taxonomy 
became  increasingly  complex.  New  forms 
were  described  in  1935,  1950,  1954,  and 
1959.  In  addition,  misidentifications  of 
Epigonus  were  published  in  several  widely 
circulated  works  on  regional  faunas  (e.g.. 
Smith,  1949b  and  1961;  Gosline  and  Brock, 
1960). 

The  aim  of  the  present  study  is  to  clarify 
the  species-level  systematics  of  the  Epi- 
gonus-\ike  fishes.  Data  from  traditional 
characters  are  evaluated  and  augmented 
by  information  from  characters  not  pre- 
viously examined  for  this  group.  A  special 
effort  is  made  to  discuss  features  such  as 
dentition  patterns  that  caused  taxonomic 
confusion  in  the  past.  The  ecology,  func- 
tional anatomy,  zoogeography,  and  evolu- 
tion of  Epigonus  will  be  discussed  in  future 
works  on  the  genus. 

METHODS 

Measurements  were  made  to  the  nearest 
tenth  of  a  millimeter  by  tlie  use  of  Helios 
needlepoint  dial  calipers;  characters  larger 
than  190  mm  were  measured  with  a  meter 
rule  or  GPM  Anthropometer.  Measure- 
ments routinely  taken  include: 

Standard  length  (SL) — from  tip  of  snout 
to  base  of  caudal  fin. 


EriaoNus  Systematics  •  Mayer       149 


Head  length  (HL) — from  tip  of  snout  to 
tip  of  opercular  spine. 

Body  depth — between  dorsal  and  v(Mitral 
surfaces  of  body  at  level  of  peKic  fin 
base. 

Head  height — from  quadratomandibular 
joint  vertically  to  bony  rim  above  eye. 

Eye  diameter — between  anterior  and  pos- 
terior margins  of  orbit  as  defined  by 
first  and  sixth  suborbitals. 

Snout  length — from  tip  of  snout  to  an- 
terior margin  of  orbit. 

Interorbital  width — shortest  distance  be- 
tween bony  rims  above  eyes. 

Maxillary  length — from  tip  of  snout  to 
posterior  margin  of  maxilla. 

Lower  jaw  length — from  tip  of  mandible 
to  quadratomandibular  joint. 

Caudal  peduncle  depth — shortest  dis- 
tance loetween  dorsal  and  ventral  sur- 
faces of  caudal  peduncle. 

Caudal  peduncle  length — from  posterior- 
most  anal  fin  ray  to  caudal  fin  base. 

First  spine  length  (first  spine  of  first 
dorsal  fin,  D,I;  first  spine  of  second 
dorsal  fin,  DJ;  second  spine  of  anal 
fin,  AH;  pelvic  fin  spine,  PJ) — from 
base  to  tip  of  spine  along  anterior  edge. 

Counts  were  made  under  a  dissecting 
microscope  with  the  use  of  dissecting 
needles  or  insect  pins.  A  Fibre-Lite  High 
Intensity  Illuminator  proved  invaluable  for 
examinations  of  oral,  branchial,  and  visceral 
structures.  Gill  raker  and  branchiostegal 
counts  were  made  on  the  left  side  of  speci- 
mens; remaining  counts  and  measurements 
were  made  on  the  right  side  whenever 
possible.  Counts  made  include:  fin  spines 
(indicated  by  Roman  numerals),  fin  rays 
(indicated  by  Arabic  numerals),  branchi- 
ostegal rays,  rakers  on  first  gill  arch,  lateral 
line  scales,  pyloric  caeca,  vertebrae  (pre- 
caudal  +  caudal,  including  hypural  fan), 
pleural  and  epipleural  ribs,  and  basal 
ptervgiophores  between  neural  spines  9 
and  10. 

Osteological  data  were  obtained  from 
radiographs  taken  at  the  Woods  Hole 
Oceanographic  Institution,  the  Museum  of 


Comparative  Zoology,  and  the  Harvard 
University  School  of  Public  Health.  Holo- 
types  of  Oxyodon  iiiacrops  and  Scepterias 
Icninwn  were  radiographed  at  the  Zoolo- 
gisches  Museum  der  Humboldt  Universitat 
and  Australian  Museum,  respectively.  More 
comprehensive  osteological  studies  were 
based  on  cleared  and  stained  specimens 
prepared  by  trypsin  digestion  (Taylor, 
1967).  Osteological  terminology  follows 
that  presented  by  Gosline  ( 1961 )  and 
Mead  and  Bradbury  (1963).  Suborbital 
bones  are  numbered  from  1  to  8  beginning 
with  the  rostralmost  element  (lacrimal). 

Statistical  data  were  analyzed  with  the 
use  of  the  Harvard  Computation  Labo- 
ratory's IBM  360/65  digital  computer. 
Standard  techniques  described  by  Mayr 
(1969:  189-193)  and  Simpson  et  al.  (1960: 
65-68,  83-88)  were  employed  for  analyzing 
meristic  data.  Morphometric  characters 
were  examined  with  the  aid  of  regression 
techniques  specified  by  Simpson  et  al. 
(1960:  215-233,  238)  and  Bailey  (1959: 
91-99). 

Before  undertaking  regression  analyses, 
morphometric  data  were  plotted  against 
SL.  Graphs  were  drawn  according  to  a 
BMD  05D  plotting  routine  (Dixon,  1967: 
71 )  and  served  as  visual  tests  for  linearity 
of  scatter.  Only  characters  exhibiting  linear 
scatters  were  analyzed  by  regression  tech- 
niques. As  a  second  precaution  against 
nonlinearity,  subadult  specimens  ( <  40 
mm  SL)  were  excluded  from  statistical 
samples. 

Data  from  several  morphometric  char- 
acters are  presented  both  as  ratios  (i.e., 
percent  of  SL  or  HL)  and  as  regression 
parameters.  The  former  are  intended  only 
as  identification  aids.  As  Royce  (1957:  17) 
points  out,  heterogenic  growth  makes  the 
use  of  ratios  in  fish  taxonomy  inefficient 
and  may  lead  to  erroneous  conclusions. 

Collection  and  institution  names  are  ab- 
breviated as  follows  in  this  paper: 

ABE        —Collection    of    Dr.    T.    Abe, 

Tokyo 
AM  — Australian    Museum,    Sydney 


150       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


BMNH  — British  Museum  (Natural 
History),  London 

BPBM  — Bernice  P.  Bishop  Museum, 
Honohihi 

CM  — Carnegie  Museum;  collections 

presently  housed  in  FMNH, 
Chicago 

DM  — Dominion  Museum,  Welling- 

ton 

EBM  — Estacion  de  Biologia  Marina, 
Universidad  de  Chile,  Viiia 
del  Mar 

FMNH  —Field  Museum  of  Natural 
History,  Chicago 

IRSN  — Institut  Royal  des  Sciences 
Naturelles  de  Belgique,  Brus- 
sels 

ISH  — Institut  fiir  Seefischerei,  Ham- 

burg 

LACM  — Los  Angeles  County  Museum 
of  Natural  History,  Los  Ange- 
les 

MCZ  — Museum  of  Camparative  Zo- 
ology, Harvard  University, 
Cambridge 

MNHN  —Museum  National  d'Histoii-e 
Naturelle,  Paris 

MZF  — Museo  Zoologico  di  Firenze, 
Florence 

RUSI  — J.L.B.  Smith  Institute  of  Ich- 
thyology, Rhodes  University, 
Grahamstown 

SAM  — South  African  Museum,  Cape 
Town 

SMF  — Natur-Museum  Senckenberg, 
Frankfurt  am  Main 

SU  — Stanford    University;    collec- 

tions presently  housed  in  the 
California  Academy  of  Sci- 
ences, San  Francisco 

TABL  — Tropical  Atlantic  Biological 
Laboratory,  Miami 

UMML  — Rosenstiel  School  of  Marine 
and  Atmospheric  Science, 
University   of   Miami,    Miami 

USNM  — National  Museum  of  Natural 
History,  Washington,  D.C. 

UZM  — Universitetets  Zoologiske  Mu- 
seum, Copenhagen 


ZMB        — Zoologisches      Museum      der 
Humboldt  Universitiit,  Berlin 

Descriptions  are  based  on  material  listed 
by  Mayer  (1972:  Appendix  II).  Additional 
data  were  obtained  from  examinations  of 
the  seventeen  specimens  listed  below.  All 
seventeen  fishes  were  radiographed. 

E.  robustus:  ISH  1132/66,  3  specimens, 
121.1-142.5  mm,  WALTHER  HER- 
WIG  Sta.  237/66,  36°00S,  52°58'W, 
800  m.  ISH  189/71,  9  specimens, 
147.0-198.0  mm,  WALTHER  HER- 
WIG  Sta.  121/71,  37°44S,  54°43'W, 
800  m.  ISH  269/71,  1  specimen,  147.5 
mm,  WALTHER  HERWIG  Sta.  340/ 
71,  38°50'S,  54°25'W,  1000  m.  ISH 
430/71,  1  specimen,  124.1  mm, 
WALTHER  HERWIG  Sta.  348/71, 
38°20'S,  54°33 W,  997-1040  m. 

E.  fmgilis:  LACM  32668-6,  1  specimen, 
72.5  mm,  2  mi.  off  Haleiwa,  Oahu, 
Hawaii,  65  fms.  SU  32262,  2  speci- 
mens, 90.0-93.9  mm,  Honolulu,  Hawaii. 

Distributions  were  determined  from 
material  examined  and  from  published 
accounts.  Because  of  the  confusion  in 
Epigomis  taxonomy,  published  data  were 
used  only  if  species  identifications  could 
be  verified  from  included  descriptions, 
illustrations,  etc.  Data  from  specimens  of 
doubtful  identity  were  not  considered.  A 
complete  list  of  station  data  taken  from  the 
literature  is  provided  by  Mayer  (1972: 
Appendix  II ) . 

No  attempt  has  been  made  to  provide 
exhaustive  synonymies  for  Epigonus  spe- 
cies. References  are  cited  only  if  they  (1) 
are  taxonomically  or  zoogeographically  im- 
portant; (2)  provide  outstanding  descrip- 
tions, illustrations,  or  synonymies;  or  (3) 
represent  verifiable  misidentifications.  Non- 
taxonomic  accounts  have  been  omitted,  as 
liave  references  to  cruise  summaries  and 
faunal  lists. 

SYSTEMATICS 

Statistical  data  are  presented  in  tables 
accompanying  species  descriptions;  meristic 


Epigonus  Systematics  •  Mayer       151 


characters  witli  low  variability  are  reported 
ill  the  text  as  value,  followed  in  parentheses 
by  number  of  specimens  exiiibiting  that 
v^alue.  Meristic  and  mensural  data  from 
holotypes  of  new  species  are  presented  in 
the  Appendix. 

Genus  Epigonus  Rafinesque,  1810 

Epifiomis  Rafinesque,  1810:  64.  (Type  .species: 
Epigonus  macrophthahnus  Rafinesque,  1810 
by  in()n()t>p\'.  A  synonym  of  Pomatomus 
telescopus  Risso,  1810.) 

Tt'h'scops  Bleeker,  1876:  261.  (Type  .species: 
Poiuatonniti  tclescopiuiu  [sic!]  Risso,  1810  by 
original  designation.  Pomatomus  deemed  in- 
applicable. ) 

Pomatomichthijs  Ciglioli,  1880:  20.  (Type  species: 
Pomatomiclithys  constanciae  Giglioli,  1880  by 
monotypy.  A  synonym  of  Pomatomus  teles- 
copus Risso,  1810.) 

Hynnodiis  Gilbert,  1905:  217.  (Type  species: 
Hijnnodus  athcrinoides  Gilbert,  1905  by  mono- 
typy. A  synonym  of  Epigonus  occidentaJis 
Goode  and  Bean,  1896.) 

Oxyodon  Brauer,  1906:  287.  (Type  species: 
Oxyodou   Diacwps  Brauer,   1906  by  monotypy.) 

Xystramia  Jordan,  1917:  46.  (Type  species: 
Glossamia  pandionis  Goode  and  Bean,  1881 
by  original  designation.  Glossamia  deemed 
inapplicable. ) 

Scepterias  Jordan  and  Jordan,  1922:  44.  (Type 
species:  Scepterias  fragilis  Jordan  and  Jordan, 
1922  by  monotypy.) 

Paraliynnodus  Barnard,  1927:  525.  (Type  species: 
Parahynnodus  robustus  Barnard,  1927  by  mono- 

typy- ) 

Diapiosis.  Epip^onus  is  distinguished 
from  other  lower  perciform  genera  by  a 
mosaic  of  characters  including  8  suborbital 
bones,  all  lacking  subocular  shelves;  large, 
thin-walled  swimbladders  with  postero- 
dorsal  ovals;  VII  or  VIII  first  dorsal  fin 
spines;  1,9  or  1,10  second  dorsal  fin  ele- 
ments; 11,9  anal  fin  elements;  15-23 
pectoral  fin  rays;  and  17-35  gill  rakers.  No 
member  of  the  genus  exhibits  fang-like 
conical  teeth,  such  as  are  found  in  Cheilo- 
dipterus,  or  anteriorly  projecting  teeth, 
such  as  are  found  in  Rosenblattia. 

Description.  Body  elongate,  fusiform; 
dorsal  and  ventral  profiles  slightly  convex, 
similar.  Mouth  oblicfue,  terminal;  upper 
jaw  protrusile;  maxilla  excluded  from  gape. 


sheathed  l)y  lacrimal  anteriorly,  free  pos- 
teriorly; supramaxilla  absent.  Eye  large, 
round  or  oval.  Nostrils  paired,  rounded  or 
slit-like,  two  on  each  side  of  head. 

Premaxillae,  mandibles,  vomer,  and  pahi- 
tines  edentulous  or  bearing  conical  teeth; 
tongue  and  endopterygoids  rarely  dc>ntiger- 
ous;  ectopterygoids  edentulous.  Gill  rakers 
moderate  to  long,  17-35;  branchiostegal 
rays  7  (3  +  4);  pseudobranchiae  present. 

Opercular  .spine  either  weak,  flattened 
and  poorly  ossified,  or  pungent  and  bony; 
spine  ventral  to  one  or  more  horny  or  mem- 
branous spinelets.  Preopercle  with  double 
edge;  angle  frequently  produced. 

Dorsal  fins  VII-1,9,  VII-1,10,  or  VIII- 
1,10,  separated  by  distinct  interdorsal 
space;  rudimentary  subcutaneous  eighth 
spine  present  in  seven-spined  forms.  Anal 
fin  11,9;  pectoral  fins  15-23;  peKic  fins  1,5: 
caudal  fin  forked,  9  +  S  principal  rays, 
upper-  and  lowermost  rays  unbranched. 

Scales  large,  deciduous,  ctenoid.  Lateral 
line  complete,  extending  parallel  to  dorsal 
profile  on  dorsolateral  surface  of  trunk, 
descending  to  midline  on  posterior  portion 
of  caudal  peduncle,  continuing  on  tail; 
lateral  line  scales  33-51;  canal  simple, 
broadening  into  deltoid  or  Y-shaped  tube 
at  rear  edge  of  scale.  Scale  pockets  cover- 
ing most  of  body  including  occiput,  soft 
dorsal,  anal,  and  caudal  fins;  scales  absent 
from  snout;  no  axillary  scale  at  base  of 
PJ  spine. 

Suborbitals  8,  all  lacking  subocular 
shelves.  Vertebrae  25;  basapophyses  on 
vertebrae  3  or  4.  Predorsals  3,  first  and 
second  interdigitating  between  neural 
spines  2  and  3,  third  located  behind  neural 
spine  3.  Caudal  skeleton  with  2  autogenous 
haemal  .spines,  6  hypurals  (hypural  1  = 
parhypural  sensu  Monod,  1968),  3  (>purals, 
2  (rarely  1)  pairs  of  uroneurals.  Actinosts 
4,  3VL'  borne  by  scapula. 

Swimbladder  large,  thin-walled,  lacking 
anterior  or  posterior  projections  to  cranium 
and  neural  arches;  diaphragm  absent;  oval 
posterodorsal;  retia  mirabilia  well  devel- 
oped.    Stomach    U-    or    Y-shaped;    pyloric 


152       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


caeca  .5-34,  may  be  modified  into  lu- 
minescent organs;  intestines  simple,  folded 
into  three  segments.  Specimens  dioecious; 
no  evidence  of  hermaphroditism  or  oral 
brooding. 

Habitat:  Engybenthic;  continental  slope 
between  approximately  200  and  1200 
meters. 

Key  to  Species  of  Epigonvs 

la  Opercular  spine  weak,  poorly  ossified,  or 
absent  (opercular  spine  refers  to  the 
ventralniost  reinforced  projection  from  the 

posterodorsal   edge   of  the   opercle)    2 

b  Opercular  spine  pungent,  bony  7 

2a  Lateral  line  scales  46-51;  tongue  eden- 
tulous or  bearing  scattered  tooth  patches 

3 

b  Lateral  line  scales  33-36;  tongue  cov- 
ered with  tooth  patches  (Fig.  lA) 
E.   oligolepis 

3a  Gill  rakers  23-34;  premaxillary  teeth  short, 
conical     or     villifonn,     not    visible     when 

mouth    closed       4 

b  Gill  rakers  17-21;  premaxillary  teetli 
elongate,  thin,  inwardly  recurved,  visible 
when  mouth  closed E.  macrops 

4a  Pyloric  caeca  7-14;  first  dorsal  fin  VII, 
rarely  VIII;  vertebral  count  10  +  15; 
specimens  not  exceeding  220  mm   SL  5 

b  Pyloric  caeca  21-34;  first  dorsal  fin  VIII, 
rarely  VII  (DiVIII  often  small  or  rudi- 
mentary); vertebral  count  11  -|-  14; 
specimens  to  550  mm  SL  E.  telescopus 

5a  Body  long,  shallow;  depth  15.8-23.6%  SL; 
peduncle    length    25.4-32.2%    SL;    caudal 

peduncle    ring    absent    6 

b  Body  short,  deep;  depth  22.4-29.6%  SL; 
peduncle  length  22.0-26.3%  SL;  caudal 
peduncle  ring  present  on  specimens 
shorter  than  110-120  mm  SL  (Fig.  IB) 
E.    pandionis 

6a  Gill  rakers  28-34;  pyloric  caeca  10-14; 
head  length  31.2-38.6%  SL;  2  pterygio- 
phores  between  neural  spines   9  and   10, 

rarely    1 E.    denticulatus 

b  Gill  rakers  25-26,  pyloric  caeca  7-8; 
head  length  30.0-34.0%  SL;  1  pterygio- 
phore  between  neural  spines  9  and  10 
E.    fiagilis 

7a  Body  moderate  to  deep,  20.0-32.0%  SL; 
dorsal    fins   VII-1,9,    rarely   VII-1,10;    gill 

rakers  26-35  8 

b  Body  shallow,  14.0-19.5%  SL;  dorsal  fins 
VII-1,10,  rarely  VII-1,9;  gill  rakers  22-27 
— .    E.    occidentalis 

8a  Gill  rakers  of  lower  arch  simple,  awl- 
shaped    9 


b  Gill   rakers  of  lower  arch  pectinate    ( Fig. 

IC)    - E.   pectinifer 

9a  Tongue    edentulous    10 

h  Tongue      covered      with      tooth      patches 

E.  trewavasac 

10a  Head  length  28.0-36.6%  SL;  head  height 
14.7-18.8%    SL;    gill    filaments    moderate 

or   short   11 

b  Head  length  36.8-41.9%  SL;  head  height 
18.9-21.1%      SL;      gill      filaments      long 

,_ E.    crassicaiidiis 

11a  Fin  spines  long,  DJ  14.8-18.7%  SL, 
All  13.0-20.8%  SL;  interorbital  width 
8.7-10.2%  SL;  eyes  large,  40.0-51.1%  HL 

E.    leinmen 

b  Fin  spines  moderate,  D2I  10.0-12.6%  SL, 
All  9.2-13.3%  SL;  interorbital  width 
6.5-8.2%  SL;  eyes  moderate  to  small, 
37.4-42.2%  HL  E.  robustus 

Epigonus  telescopus  (Risso,  1810) 
Figure  2 

Pomatomus  telescopus  Risso,  1810:  301,  plate  IX, 
fig.  31  (original  description;  Nice;  holotype 
examined,  MNHN  B862);  Lowe,  1841:  173; 
Capello,  1868:  160;  Moreau,  1881:  386,  fig. 
125;  Vaillant  (in  part),  1888:  376. 

Epigonus  macwphthalmus  Rafinesque,  1810:  64 
( original  description;  no  type  locality;  holotype 
lost). 

Pomatomus  telescopium  Cuvier,  1828:  171  (in- 
correct emendation  of  Pomatomus  telescopus 
Risso,  1810);  Valenciennes,  1830:  495; 
Valenciennes,  1837-1844:  6,  plate  I;  Giinther, 
1859:  250;  Cocco,  1885:  85;  Holt  and  Calder- 
wood,  1895:  405,  plate  LXIl. 

Pomatomus  cuvieri  Cocco,  1829:  143  (original 
description;  seas  of  Messina;  holotype  not 
examined ) . 

Pomatotnus  cuvicrii  \'alenciennes,  1830:  501  (in- 
correct emendation  of  Pomatomus  cuvieri 
Cocco,  1829). 

?Pomatomichthys  constanciae  Giglioli,  1880:  20 
( original  description;  Straits  of  Messina;  holo- 
type not  examined,  MZF  3089);  Goode  and 
Bean,   1896:   234. 

Epigonus  telescopus  Goode  and  Bean,  1896:  232; 
Cligny,  1903:  9;  Barnard,  1927:  523;  Gall, 
1931:  1,  fig.  1;  Fowler,  1936:  736,  fig.  326; 
Smith,  1949b:  206,  fig.  474. 

Scepterias  lenimen,  Whitley  ( in  part )  ( not  Whit- 
ley, 1935),  1968:  56. 

Diagnosis.  E.  telescopus  is  the  largest 
species  of  the  genus,  growing  to  over  550 
mm  SL.  Specimens  are  characterized  by 
21-34  pyloric  caeca  and  eight  first  dorsal 


EriaoNvs  Systematics  •  Mayer       153 


B 


j^i^' 


^^.^fffi^^'^h'"**'^^' 


Figure  1.  A.  Tongue  of  E.  oligolepis.  Stippled  areas 
indicate  tooth  patches;  shape  and  size  of  tooth  patches 
may  vary  among  specimens.  B.  Caudal  peduncle  of 
young  E.  pandionis  showing  anterior  ring  and  posterior 
band.  C.  Gill  raker  of  E.  pectinifer  showing  nub-like 
processes. 


fin  spines.  The  opercular  spine  is  blunt  and 
poorly  ossified  and  distinguishes  the  species 
from  E.  occidentalis,  E.  trewavosae,  E. 
pectinifer,  E.  rohiistus,  E.  lenimen,  and  E. 
crassicaudus,    which    have   pungent    oper- 


Y12yc 


cular  spines.  Unlike  remaining  congeners, 
E.  telescopus  possesses  11  +  14  vertebrae. 

Description.  Meristic  data  presented  in 
Table  1;  regression  data  for  morpho metric 
traits  presented  in  Table  2. 

Body  thickset,  shortened;  anterodorsal 
profile  slightly  convex,  rising  most  steeply 
from  tip  of  snout  to  interorbital  region; 
body  moderate  to  deep,  21.2-26.3%^  SL; 
caudal  peduncle  short,  19.0-26.5%  SL. 

Head  moderate  to  deep,  height  13.3- 
SL;  length  30..5-37.9%  SL;  snout 
blunt;  angle  of  gape  moderate  to  large; 
lower  jaw  equalling  or  protruding  slightly 
beyond  upper  jaw.  Maxilla  rarely  exceed- 
ing %-%  eye  length,  posterior  margin  of 
maxilla  broad,  posteriormost  point  near 
ventral  surface  of  bone;  maxillae  of  large 
specimens  scaled.  Eye  round,  49.5-58.9% 
HL;  circumorbital  tissues  scaled,  scale 
pockets  particularly  apparent  in  large  spec- 
imens; anterodorsal  rim  of  orbit  projecting 
into  profile  in  small  forms,  reaching  profile 
in  larger  forms;  interorbital  width  9.0- 
10.9%  SL. 

Dentition  variable  with  age  (see  Onto- 
genetic change);  premaxillae,  mandibles, 
vomer,  and  palatines  dentigerous;  tongue 
edentulous. 

Opercle  bearing  short,  poorly  ossified 
spine  ventral  to  1-8  membranous  or  poorly 
ossified  spinelets;  spine  and  spinelets  sepa- 
rated by  shallow  gap;  spinelets  occasionally 
obscured  by  underlying  membranes.  Pre- 
opercle  variable  with  age;  angle  rounded, 
slightly  produced  in  specimens  shorter  than 


Figure  2.   Epigonus  telescopus,  220.0  mm  SL,  ISH  70/63. 


154       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  1.     Epigonus  telescopus  meristic  data.  X  =  mean;  SD  =  stan- 
dard DEVIATION;  n  =  NUMBER  OF  SPECIMENS. 


Range 


SD 


Pectoral    fin    rays  20.85  19-23  0.71  54 

Gill    rakers  24.40  23-26  0.85  52 

Lateral    line    scales  48.14  46-50  1.09  50 

Pyloric    caeca  25.25  21-34  3.59  16 


200  mm  SL,  broadly  produced  in  larger 
forms;  minute  serrations  on  angle  and  ven- 
tral surface  of  bone,  rarely  along  posterior 
surface  dorsal  to  angle;  striations  radiating 
from  inner  edge  of  angle.  Interopercles 
and  subopercles  without  stiiations,  occa- 
sionally bearing  minute  serrations  on  pos- 
tero ventral  surfaces.  Gill  rakers  simple, 
awl-like. 


First  dorsal  fin  VII  (7),  VIII  (46), 
eighth  spine  small  or  rudimentary,  lack- 
ing membranous  connection  to  preceding 
spines;  second  dorsal  fin  1,9  (1),  1,10  (52), 
1,11  (1);  anal  fin  11,9  (56);  D,I  long, 
3.5-6.3%  SL;  DJ,  All  short,  5.3-9.5%, 
5.7-10.6%o  SL  respectively;  Pol  moderate, 
6.5-11.9%  SL. 

Vertebrae  11  +  14  (18);  epipleural  ribs 


Table  2.    Epigonus  telescopus  regression  data,  b  =  regression  coeffi- 
cient   ±    95%o    CONFIDENCE   INTERVAL;    a    =    Y   INTERCEPT;    n    =    NUMBER   OF 

specimens.   All  regressions  on  SL. 


b 

a 

n 

HL 

0. 

35 

+ 

0. 

01 

1. 

60 

50 

Body    depth 

0. 

25 

+ 

0. 

01 

-2, 

43 

48 

Head    height 

0. 

19 

+ 

0. 

00 

0. 

45 

45 

Eye    diameter 

0. 

13 

+ 

0. 

01 

6. 

57 

44 

Snout    length 

0. 

10 

+ 

0. 

00 

-2. 

19 

49 

Interorbital    width 

0. 

10 

+ 

0. 

00 

-0, 

32 

52 

Maxillary    length 

0. 

16 

+ 

0. 

00 

0. 

00 

48 

Lower    jaw    length 

0. 

19 

+ 

0. 

00 

-0. 

61 

50 

Caudal   peduncle    de 

pth 

NONLINEAR 

Caudal  peduncle    length 

0. 

2  1 

+ 

0. 

01 

3. 

35 

51 

D2I 

0. 

06 

+ 

0. 

02 

3. 

89 

1  1 

All 

0. 

06 

+ 

0. 

01 

4. 

18 

31 

P2I 

NONLINEAR 

Ei'iaoNus  Systematics  •  Mayer       155 


Table  3.     Ontogenetic  changes  in  the  dentition  of  E.  telescopus. 


A.    PREMAXILLARY     DENTITION 

<  200   mm   SL 

Extent  1/2-2/3  of  ventral 

surfoce 


Pottern 


I    row 


200-400  mm  SL 

2/3-7/8  of  ventrol 
surface 


1-2  irregular   rows 
tapering  to  I    row 


>  400  mm  SL 

2/3-7/8  of  ventral 
surface 

Multiple    irregular  rows 


B.    MANDIBULAR    DENTITION 


Extent 


<  150  mm  SL 
Entire  coronoid  surface 


150-250    mm  SL 
Entire  coronoid  surface 


>  250  mm  SL 
Entire  coronoid  surface 


Pottern 


row 


2-3  irregular  rows 
tapering  to  |-2  rows 


3, 4,  or  5    irregular  rows 


C.    VOMERINE    DENTITION 


Extent 


<    I  75  mm  SL 
Center  of  vomer 


>   I  75  mm   SL 
Entire    face  of  vomer 


Pattern 


Scattered  teeth  in  few 
irregular  rows 


Numerous  teeth    in    multiple 
irregular  rows 


D.    PALATINE     DENTITION 


Extent 


<   150  mm  SL 
Length  of  ventral   surface 


>  I  50  mm   SL 
Length   of   ventral  surface 


Pottern 


1-2   irregular  rows 
tapering  to  I  row 


2-5  irregular  rows 
tapering  to  I  row 


7  (11),  8  (2),  inserting  on  vertebrae  1-7 
or  1-8  respectively;  pleural  ribs  9  (17), 
inserting  on  vertebrae  3-11. 

Large  specimens  black  or  brown-violet, 
iridescent  in  life  ( Risso,  1810;  Steindachner, 
1891;  Dons,  1938).  Color  in  alcohol  vari- 
able with  mode  of  collection  and  preser- 
vation; skin  often  abraded,  revealing  under- 
lying white-orange  tissue;  scale  pockets 
mottled  with  black  or  brown,  melanophores 
more  densely  packed  near  caudal  edges; 
pigment  darker  in  larger  fish;  skin  oily, 
cutaneous  fat  deposits  adding  rust-colored 
tint;  opercular  area  black.  Guanine  de- 
posits occasionally  occurring  on  opercular, 
tlioracic,  and  abdominal  regions;  iris  black 


with  silver  highlights;  branchial  membranes 
black;  mouth  darkening  with  age  (see 
Ontogenetic  change ) . 

Description  based  on  54  specimens  68.1- 
553  mm  SL. 

Ontogenetic  change.  Several  marked 
ontogenetic  changes  occur  in  E.  telescopus, 
the  most  noticeable  involving  dentition 
patterns.  Tooth-bearing  bones  of  young 
specimens  exhibit  relatively  prominent  con- 
ical teeth.  Teeth  become  more  numerous 
witli  growth  but  appear  smaller  and  form 
weak  conical  or  villiform  bands.  As  Table 
3  illustrates,  older  specimens  have  more 
complex  tooth  patches  with  larger  numbers 
of  tooth  rows. 


156       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Epigonus  Systematics  •  Mayer       157 


A  second  change  involves  oral  pigmenta- 
tion. Young  individuals  have  white  or  pale 
yellow  mouths;  melanin  is  present  only  in 
the  vicinity  of  the  pharynx.  By  the  time 
specimens  reach  175-225  mm  SL,  black 
pigment  extends  anteriorly  to  cover  the 
entire  tongue.  Shortly  thereafter,  the  palate 
becomes  totally  blackened,  and  by  300 
mm  SL,  the  entire  mouth  is  dark. 

The  above  changes  are  associated  with 
alterations  in  intestinal  length.  Measure- 
ments of  fourteen  specimens  ranging  from 
90.7-553  mm  SL  indicate  that  intestinal 
length  increases  from  66-737^  SL  in  small 
specimens  (90.7-128.5  mm)  to  98-108% 
SL  in  moderate-sized  individuals  (220-250 
mm).  Thereafter,  intestines  grow  more 
slowly,  reaching  110-115%  SL  in  the  largest 
specimens.  The  coincidence  of  rapid  in- 
testinal growth,  dentition  changes,  and  de- 
velopment of  oral  pigment  suggests  that 
E.  telescopus  modifies  its  feeding  habits 
with  growth. 

Distribution.  E.  telescopus  has  an  anti- 
tropical  distribution  in  the  Atlantic,  oc- 
curring from  Iceland  to  the  Canary  Lslands 
and  reappearing  along  the  western  coast 
of  South  Africa  (Fig.  3).  Specimens  have 
also  been  taken  in  the  Subtropical  Con- 
vergence region  east  of  New  Zealand.  The 
species  is  well  known  in  the  western  Med- 
iterranean and  has  been  captured  once 
off  the  eastern  coast  of  North  America.  A 
single  specimen  is  known  from  shallow 
water  off  Norway  (Dons,  1938). 

Adults  are  taken  by  bottom  trawl  or  long- 
line  and  are  most  abundant  from  300  to 
800  meters;  however,  specimens  have  been 
captured  from  water  as  shallow  as  75  to 
80  meters  to  as  deep  as  1000  to  1200  meters. 
Koefoed  (1952)  reports  four  pelagic  ju- 
veniles from  the  Azores;  Bertolini  (1933) 
mentions  the  presence  of  juveniles  in  the 
Tyrrhenian  Sea. 

Earlier  workers  reported  the  range  of  E. 
telescopus  to  include  St.  Helena  (Val- 
enciennes, 1837-1844;  Giinther,  1868; 
Bauchot  and  Blanc,  1961),  tropical  west 
Africa   (Osorio,  1898;  Poll,  1954;  Bauchot 


and  Blanc,  1961),  and  the  Indian  Ocean 
(Steindachner,  1907;  Fowler,  1935).  These 
accounts  are  based  on  misidentified  or 
tenuously  identified  material.  The  .speci- 
mens described  by  Giinther,  Poll,  and 
Bauchot  and  Blanc  are  E.  pandionis,  while 
that  examined  by  Fowler  is  Scomhrops- 
like.  Valenciennes'  identification  is  ba.sed 
on  an  impublished  description  and  figure 
by  a  St.  Helena  resident  and  must  be  re- 
garded with  suspicion.  Reports  by  Stein- 
dachner and  Osorio  could  not  be  evaluated, 
because  neither  includes  a  description  or 
figure  of  the  material  studied. 

Geographic  variation.  The  scarcity  of 
material  from  South  Africa  and  New 
Zealand  makes  it  difficult  to  judge  the 
degree  to  which  Northern  and  Southern 
Hemisphere  populations  of  E.  telescopus 
have  diverged.  Comparisons  of  dorsal  and 
pectoral  fins,  pleural  and  epipleural  ribs, 
lateral  line  scale  counts,  gill  rakers,  and 
pyloric  caeca  reveal  no  subspecific  dif- 
ferences (coefficients  of  difference  ^  0.44). 
Moiphometric  characters,  on  the  other 
hand,  exhibit  greater  variability.  Of  eight 
traits  successfully  analyzed,  three  are  sig- 
nificantly different  at  both  the  95%,  98%, 
and  99%  levels  of  confidence  (Table  4). 
These  differences  suggest  that  northern 
and  southern  populations  represent  gemi- 
nate subspecies;  however,  additional  ma- 
terial must  be  collected,  especially  from  the 
Southern  Hemisphere,  before  definitive 
statements  can  be  made  on  intraspecific 
variability. 

Ta.xonomic  notes.  Pomatomichthtjs  con- 
stanciae  Giglioli,  1880  is  pro\'isionaily  con- 
sidered a  junior  synonym  of  E.  telescopus 
on  the  basis  of  work  by  Tortonese  and 
Queirolo  (1970).  These  authors  re-exam- 
ined and,  for  the  first  time,  figured  the 
holotype  of  P.  constanciae.  The  latter 
species  is  known  only  from  the  type  speci- 
men. The  original  description  (Giglioli, 
1880)  is  incomplete;  no  adequate  rede- 
scription  has  ever  been  published. 

Data  from  the  papers  mentioned  above 
indicate  a  similarit\'  between  P.  constanciae 


158       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  4.  Comparison  of  regression  coefficients  from  Northern  and  Southern 
Hemisphere  populations  of  E.  telescopus.  Data  evaluated  at  the  95%,  98%, 
and  99%  levels  of  confidence.  df  =  degrees  of  freedom;  nb  =  regression 
coefficients  of  Northern  Hemisphere  specimens;  Sb  =  regression  coefficients 
OF  Southern  Hemisphere  specimens;  SD  =  significant  difference  between  tab- 
ular AND  calculated  VALUES  OF  t;  t  ^  CALCtJLATED  VALUES  OF  t. 

Significance 


irt 

CO 

05 

Nb 

Sb 

DF 

t 

a> 

CT> 

05 

HL 

0. 

35 

0 

36 

46 

1.  54 

Body    depth 

0. 

25 

0 

24 

44 

0.  82 

Head    height 

0. 

20 

0 

18 

41 

3.  88 

SD 

SD 

SD 

Eye    diameter 

0. 

13 

0 

12 

40 

1.  75 

Snout    length 

0. 

09 

0 

10 

45 

3.  19 

SD 

SD 

SD 

Interorbital    width 

0. 

10 

0 

10 

48 

1.  05 

Maxillary    length 

0 

15 

0 

.  16 

12 

0  .  16 

Lower    jaw    len 

gth 

0 

19 

0 

.  19 

14 

0.  71 

Caudal    pedunc 
depth 

le 

NONLINEAR 

Caudal    pedunc 
length 

le 

0 

21 

0 

.  22 

47 

0.  58 

^2^ 

INSUFFICIENT 

DATA 

All 

0 

06 

0 

.  04 

23 

3.  80 

SD 

SD 

SD 

^2^ 

NONLINEAR 

Table  5.     Comparison  of  dorsal  and  pectoral  fin  counts  from  E.  telescopus,  P. 

CONSTANCIAE,    AND    E.    TREWAVASAE.      DaTA    FOR    P.    CONSTANCIAE    FROM    GiGLIOLI     (1880) 
AND   TORTONESE   AND   QuEIROLO    (1970);    REMAINING   DATA    FROM    PRESENT    STUDY. 

E.     telescopus  P.     constanciae  E.     trewavasae 


VII  VII 

I,  9. 
1,9  rarely 

I,  10 

18  16-18 


First 

VIII, 

dorsal 

rarely 

fin 

VII 

Second 

I,  10, 

dors  a 1 

r  ar ely 

fin 

I,  9 

Pectoral 
fin 

19  -  2: 

Ei'iGONus  Systematics  •  Mayer       159 


Figure  4.     Epigonus  macrops,  154.6  mm  SL,  USNM  207679. 


and  E.  telescopus  but  also  suggest  an  af- 
finity between  P.  constanciae  and  E.  tre- 
wavasae  Poll,  1954.  As  is  shown  in  Table 
5,  dorsal  and  pectoral  fin  counts  fall  within 
the  range  of  E.  treicavasae  rather  than  E. 
telescopus.  Tortonese  and  Queirolo's  figure 
similarly  shows  the  holotype  to  possess  a 
sharp  opercular  spine,  short  DJ,  and  long 
PJ — all  characteristics  of  E.  treicavasae. 
Mensural  data  fail  to  differentiate  P. 
constanciae  from  either  species.  Unlike  E. 
treicavasae  but  like  E.  telescopus,  the  holo- 
type lacks  lingual  teeth  (Giglioli,  1880). 

In  view  of  the  uncertainty  surrounding 
P.  constanciae,  a  closer  study  of  this  form 
must  be  undertaken.  The  problem  is  all 
the  more  pressing,  because  E.  treicavasae 
is  recorded  from  the  Mediterranean  for  the 
first  time  in  this  paper. 

Common  names.  Comprehensive  lists  of 
common  names  for  E.  telescopus  are  pro- 
vided by  Doderlein  (1889),  Nobre  (1935), 
and  Bini  (1968).  Three  names  not  re- 
corded in  these  works  are  "Mejluza" — Gran 
Canaria  ( Steindachner,  1891),  "Devil-fish" 
— North  Sea  area  (Ehrenbaum,  1928),  and 
"Big-eyed  cardinal  fish" — New  Zealand 
( Anonymous,  1961 ) . 

Epigonus  macrops  (Brauer,  1906) 
Figure  4 

Oxijdon  macrops  Brauer,  1906:  288,  fig.  172 
(original  description;  Indian  Ocean,  land-locked 
sea  on  west  coast  of  Sumatra,  VALDIVIA  Sta. 
186,  03°21'01"S,  101°11'05"E,  903  m;  syntype 
examined,  ZMB  17678);  Weber  and  de  Beau- 
fort, 1929:  351,  fig.  81;  Nomian,  1939:  60. 


Diagnosis.  E.  macrops  may  be  distin- 
guished from  all  congeners  by  its  low  gill 
raker  counts  ( 17-21 ) .  It  is  further  char- 
acterized by  eight  fully  developed  first 
dorsal  fin  spines  and  eight  pyloric  caeca, 
one  of  which  may  function  as  a  lumin- 
escent organ. 

Description.  Meristic  values  presented 
in  Table  6;  regression  data  for  morpho- 
metric  traits  presented  in  Table  7. 

Body  elongate;  anterodorsal  profile  rising 
steeply  to  occipital  area;  thereafter,  weakly 
convex,  almost  horizontal  to  first  dorsal  fin; 
body  depth  19.7-24.1%  SL;  caudal  pe- 
duncle length  22.0-26.7%  SL. 

Head  length  34.1-38.5%  SL;  head  licight 
17.2-21.9%  SL;  snout  blunt;  angle  of  gape 
large;  lower  jaw  protruding  beyond  upper 
jaw.  Maxilla  rarely  exceeding  Vs-%  eye 
length;  posterior  margin  of  maxilla  broad, 
bearing  posteriormost  point  at  ventral  sur- 
face of  bone.  Eye  round  to  oval,  39.7- 
48.3%'  HL;  anterodorsal  rim  of  orbit  pro- 
jecting strongly  into  dorsal  profile;  inter- 
orbital  region  wide,  9.5-11.7%  SL. 

Teeth  conical,  frequently  recurved.  Pre- 
maxillary  and  mandibular  teeth  prominent, 
needle-like,  arranged  in  single  row  along 
length  of  jaws;  mandibular  teeth  occa- 
sionally forming  double  row  near  sym- 
physis; vomerine  teeth  few,  moderate, 
arranged  in  2-4  irregular  rows  or  in  a 
triangular  or  diamond-shaped  patch;  pala- 
tin(\s  bearing  2-6  teeth,  arranged  in  single 
row  covering  anterior  half  or  second  quar- 
ter of  bone;  tongue  edentulous. 

Opercular  spine  short,  weak,  bony,  ven- 


160       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  6.     Epigonus  macrops  mebistic  data.  X  =  mean;  SD  =  standard 

DEVIATION;   n   =  NLTNIBER  OF  SPECIMENS. 


X 


Range 


SD 


Pectoral    fin    rays  18.87  18-19  0.35  30 

Gill    rakers  18.63  17-21  0.87  32 

Lateral    line    scales  48.61  46-50  0.83  28 

Pyloric    caeca  8.00  8  0.00  15 


tral  to  3-10  spinelets;  spine  and  spinelets 
separated  by  shallow,  occasionally  narrow 
gap.  Preopercular  angle  weakly  produced, 
rounded,  serrate;  serrations  occasionally  ex- 
tending to  posterior  and  ventral  surfaces  of 
bone,  rarely  absent;  striations  radiating 
from  inner  edge  of  angle.  Subopercle  and 
interopercle  generally  serrated,  occasionally 
striated.   Gill  rakers  short,  awl-like. 


First  dorsal  fin  VII  (1),  VIII  (29); 
second  dorsal  fin  1,9  (1),  1,10  (31);  anal 
fin  11,9  (30),  11,10  (1).  D^I,  DJ,  All 
short,  equalhng  1.2-2.9%,  5.3-7.7%,  5.9- 
9.9%  SL  respectively;  PJ  moderate,  11.7- 
14.1%  SL. 

Vertebrae  10  +  15  (25);  epipleural  ribs 
6  (23),  inserting  on  vertebrae  1-6;  pleural 
ribs  8  (24),  inserting  on  vertebrae  3-10. 


Table  7.     Epigonus  macrops  regression  data,    b  =  regression  coeffi- 
cient ±  95%  confidence  interval;  a  =  Y  intercept;  n  =:  number  of 
specimens.   All  regressions  on  SL. 


b 

a 

n 

HL 

0. 

35 

+ 

0. 

02 

1 

±  • 

48 

26 

Body    depth 

0, 

22 

+ 

0. 

02 

0. 

51 

31 

Head    height 

0. 

18 

+ 

0. 

02 

1. 

74 

19 

Eye    diameter 

0, 

14 

+ 

0. 

02 

3. 

41 

29 

Snout    length 

0. 

08 

+ 

0. 

01 

-0. 

18 

22 

Interorbital    wi 

dth 

0, 

11 

+ 

0. 

01 

0. 

36 

30 

Maxillary    leng 

th 

0. 

14 

+ 

0. 

01 

1. 

31 

22 

Lower    jaw    leu 

gth 

0, 

18 

+ 

0. 

01 

1. 

18 

31 

Caudal    peduncl 

e    dep 

th 

0. 

12 

+ 

0. 

01 

-  1. 

53 

29 

Caudal    peduncl 

e    len 

gth 

0. 

24 

+ 

0. 

02 

0. 

70 

30 

D2I 

0. 

02 

+ 

0. 

02 

7. 

05 

13 

All 

0. 

04 

+ 

0. 

01 

4. 

73 

22 

P2I 

0. 

13 

+ 

0. 

02 

0. 

49 

16 

Ei'icoNus  Systematics  •  Mayer 


161 


Figure  5.     Caudal  peduncle  of  young  E.  macrops  bear- 
ing anterodorsally  canted  ring. 


Specimens  probably  black  in  life.  Color 
in  alcohol  variable  with  preservation;  scale 
pockets  covered  with  black  melanophores 
near  posterior  edges;  skin  trecjiiently 
abraded,  revealing  pink-yellow  muscnla- 
ture;  opercular  bones  transparent,  colored 
black  by  underlying  branchial  membranes; 
iris  black;  mouth  black  in  adults.  Young 
bearing  anterodorsally  canted  caudal  pe- 
duncle ring  (see  Ontogenetic  change). 
First  pyloric  caecum  modified  into  lumin- 
escent organ  ( see  Remarks ) . 

Description  based  on  32  specimens  77.8- 
206.0  mm  SL. 

Ontogenetic  change.  The  transition  from 
juvenile  to  adult  in  E.  macrops  is  marked 
by  changes  in  pigmentation  and  body 
shape.  Pelagic  juveniles  15-37.9  mm  SL 
and  young  demersal  forms  77.8-79.8  mm 
SL  bear  a  thin,  black,  anterodorsally  tilted 
ring  circling  the  center  portion  of  the 
caudal  peduncle  (Fig.  5).  Specimens 
larger  than  90  mm  SL  lack  this  marking. 
Melanophores  forming  the  rings  are  deeply 
embedded  in  the  peduncle  musculature 
and  cannot  be  obliterated  by  abrading  the 
surface  of  the  fish. 

Adult  E.  macrops  arc  characterized  by 
black  oral  and  branchial  membranes.  Al- 
though these  areas  are  colorless  or  poorly 
pigmented  in  specimens  smaller  than  40 
mm  SL,  the  former  surfaces  darken  and  the 
latter  become  covered  with  brown  melano- 
phores by  the  time  fish  reach  80  nun  SL. 

Juvenile  E.  macrops  appear  longer 
and  shallower  than  adults.  Ratio-on-size 
diagrams  for  interorbital  width  (i.e.,  in- 
terorbital  width/ SL  vs.  SL)  indicate  al- 
lometric  growth  takes  place  in  small 
specimens.   Similar  statements  are  probably- 


true  for  head  height,  eye  length,  and 
caudal  peduncle  measurements  but  could 
not  be  tested  because  of  damage  to  juvenile 
specimens. 

DistriJnition.  E.  macrops  adults  are  taken 
exclusively  by  bottom  trawls  between  550 
and  1100  meters  in  the  Lidian  Ocean, 
Gulf  of  Mexico,  Caribbean  Sea,  and  West- 
ern Atlantic.  Specimens  are  most  abundant 
between  640  and  920  meters.  Pelagic  ju- 
veniles are  known  from  the  Caribbean  at 
depths  of  120  to  550  meters  (Fig.  6). 

GcograpJiic  variation.  No  investigation 
made  because  of  inadec^uate  Indian  Ocean 
samples. 

Taxonomic  notes.  Brauer's  description  of 
Oxyodon  macrops  ( 1906 )  is  based  on  two 
syntypes  from  the  eastern  Indian  Ocean 
( 172  and  212  mm  total  length ) .  Of  these, 
only  the  larger  is  in  the  Zoologisches  Mu- 
seum der  Humboldt  Universitiit;  the  smal- 
ler has  been  lost.  The  misplaced  type  may 
have  been  deposited  in  the  Zoologisches 
Institut  der  Universitiit  Leipzig  and  may 
reappear  when  portions  of  this  collection, 
presently  stored  in  Berlin,  are  sorted  and 
catalogued  (Karrer,  personal  communica- 
tion ) . 

Remarks.  Specimens  of  E.  macrops  bear 
eight  pyloric  caeca;  one  of  these  appears 
modified  into  a  bioluminescent  organ.  The 
luminescent  caecum  arises  from  the  mid- 
ventral  surface  of  the  pylorus  just  before 
the  duodenum  and  main  body  of  pyloric 
appendages  (Fig.  7).  It  extends  ventrally 
until  it  reaches  the  floor  of  the  abdominal 
cavity,  bends  anteriorly  and  inserts  into  a 
pouch  formed  by  the  black  peritoneal 
lining  of  the  body  cavity.  At  the  posterior 
edge  of  the  pelvic  girdle,  the  caecal  pouch 
lies  over  a  thin,  translucent  portion  of  the 
body  wall  which  may  function  as  a  biolu- 
minescent window.  Externally  the  biolumi- 
nescent window  is  covered  by  a  single 
large  scale.  The  caecal  pouch  is  lined  with 
silver  or  silver-gray  pigment.  Guanine 
deposits  appear  most  concentrated  anter- 
odorsally. 

.\lthouah  there  is  no  direct  evidence  to 


162       Bulletin  Museum  of  Coiiiparative  Zoology,  Vol.  146,  No.  3 


m 
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Epiconus  Systematics  •  Mayer       163 


Figure  7.     Luminescent  organ  of  E.  macrops.    BW,  body  wall;  D,  duodenum;  LPC,  luminescent  pyloric  caecum; 
LW,  luminescent  window;  PC,  nonluminescent  pyloric  caeca;  PER,  peritoneum;  R,  reflector;  S,  stomach. 


support  tlie  claim  that  E.  macrops  is  lu- 
minescent, the  modifications  described 
above  are  similar  to  those  found  in  several 
luminescent  perciforms.  Pernpheris  klun- 
zingeri  and  Parapriacanthus  ransonneti 
(Pempheridae)  have  luminescent  organs 
embedded  in  the  thoracic  ventral  muscula- 
ture formed  from,  or  directly  associated 
with,  the  first  pair  of  pyloric  caeca  ( Haneda 
et  al.,  1966).  Luminescent  shallow-water 
apog(jnids  such  as  Apogon  ellioti  and 
Siphamia  nwiirnai  also  have  luminescent 
organs  associated  with  the  alimentary  canal. 
In  both  of  the  latter  forms,   anal  and/ or 


thoracic  organs  are  connected  by  duct  to 
the  intestine.  As  in  E.  nuicrops,  tissue 
above  the  luminescent  structures  may  serve 
as  a  reflector  (Iwai,  1959;  Haneda  et  al., 
1966). 

Common  names.  None. 

Epigonus  pandionis  (Goode 
and  Bean,  1881) 
Figure  8 

Apogon  pandionis  Goode  and  Bean,  1881:  160 
( original  description;  off  entrance  to  Cliesa- 
peake  Bay;  holotype  examined,  USNM  26228); 
Jordan  and  Gilbert,  1882:  564. 


Figure  8.     Epigonus  pandionis,  141.7  mm  SL,  TABL  uncatalogued. 


164       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  8.     Epigonus  pandionis  meristic  data.    X  =  mean;  SD  =  stan- 
dard deviation;  n  =  number  of  specimens. 


X 


Range 


SD 


Pectoral    fin    r  ays 
Gill    rakers 
Lateral    line    scales 
Pyloric    caeca 


17.  81 

17-  19 

0.  57 

97 

27.  84 

26-  30 

0.  88 

101 

47.  63 

46-  49 

0.  66 

81 

10.  81 

10-  13 

0.  74 

72 

Glossamia  pandionis  Goode  and  Bean,  1896:  231. 

Xystramia  pandionis  Jordan,  1917:  46. 

Epigonus    telescopus.     Poll     (not     Risso,     1810), 

1954:  89,  fig.  26;  Bauchot  and  Blanc  (in  part), 

1961:  70. 

Diagnosis.  E.  pandionis  is  the  most  ro- 
bust species  of  the  genus.  Specimens 
shorter  than  110-125  mm  SL  are  dis- 
tinguished by  a  posterodorsally  canted  ring 
circhng  the  caudal  peduncle. 

E.  pandionis  differs  from  E.  macrops  and 
E.  telescopus  by  bearing  VII  (rarely  VIII) 
spines  in  the  first  dorsal  fin  and  10-13 
pyloric  caeca.  It  is  unlike  E.  oligolepis 
because  it  has  46-49  lateral  line  scales  and 
may  be  distinguished  from  E.  treioavasae, 
E.  pectinifer,  E.  robustus,  E.  lenimen,  E. 
crassicaudus,  and  E.  occidentalis  because 
it  lacks  a  pungent,  bony  opercular  spine. 
E.  pandionis  most  closely  resembles  E. 
fragilis  and  E.  denticulatus  but  is  differ- 
entiated by  its  short  caudal  peduncle  ( 22.0- 
26.3%  SL)  and  deep  body  (22.4-29.6% 
SL).  It  further  differs  from  E.  dejiticulatiis 
by  exhibiting  gill  raker  counts  of  26—30 
and  a  single  basal  pterygiophore  between 
neural  spines  9  and  10. 

Description.  Meristic  values  presented  in 
Table  8;  regression  data  for  morphometric 
traits  presented  in  Table  9. 

Body  shortened,  robust;  anterodorsal  pro- 
file convex,  particularly  between  occiput 
and  first  dorsal  fin;  body  deep,  22.4-29.6% 
SL;  caudal  peduncle  short,  broad,  length 
22.0-26.3%  SL. 

Head  length  33.0-39.0%  SL;  head  height 


19.0-22.2%  SL;  snout  blunt;  angle  of  gape 
large;  upper  jaw  subequal  to  lower  jaw. 
Maxilla  reaching  %-y-2  eye  length;  pos- 
terior margin  broad,  posteriormost  point  at 
ventral  edge  of  bone.  Eye  round  or  slightly 
oval,  37.4-48.7%  HL;  anterodorsal  rim  of 
orbit  projecting  into  profile  in  smaller 
specimens,  reaching  profile  in  larger  forms; 
interorbital  region  wide,  9.2-11.5%  SL. 

Dentition  variable  with  age  (see  Onto- 
genetic change);  premaxillae,  mandibles, 
vomer,  and  palatines  dentigerous,  bearing 
conical,  occasionally  recurved  teeth;  tongue 
edentulous. 

Opercular  spine  short,  horny,  ventral  to 
2-5  (usually  3-4)  poorly  ossified  spinelets; 
spine  separated  from  spinelets  by  gap; 
spinelets  occasionally  obscured  by  un- 
derlying membranes.  Preopercular  angle 
broad,  rounded,  moderately  produced; 
striations  radiating  from  inner  ridge  to 
edges  of  angle;  serrations  along  posterior 
and  ventral  surfaces  of  bone.  Subopercle 
and  interopercle  bearing  scattered  serra- 
tions. Gill  rakers  awl-like. 

First  dorsal  fin  VII  (95),  VIII  (5);  sec- 
ond dorsal  fin  1,9  (2),  1,10  (97),  1,11  (1), 
11,10  (1);  anal  fin  11,8  (2),  11,9  (96),  11,10 
(1);  DJ  long,  3.6-8.1%  SL;  DJ,  All,  PJ 
short,  equalling  5.0-8.6%,  5.0-8.7%,  8.9- 
12.7%  SL  respectively. 

Vertebrae  10  +  15  (36);  epipleural  ribs 
6  ( 25 ) ,  7  ( 1 ) ,  inserting  on  vertebrae  1-6 
and  1-7  respectively;  pleural  ribs  8  (36), 
inserting  on  vertebrae  3-10. 


Epigonus  Systematics  •  Mayer       165 


Table  9.     Epigonus  pandionis  regression  data,  b  =  regression  coefft- 

CIENT    ±    95%    CONFIDENCE    INTERVAL;    a    =    Y    INTERCEIT;    11    =    NUMBER   OF 

SPECIMENS.    All  regressions  ON  SL. 


b 

a 

n 

HL 

0. 

36 

+ 

0. 

01 

0. 

04 

77 

Body    depth 

0. 

29 

+ 

0. 

01 

-2. 

64 

75 

Head    height 

0. 

21 

+ 

0. 

01 

-0. 

30 

67 

Eye    diameter 

0. 

16 

+ 

0. 

01 

-0. 

15 

80 

Snout    length 

0. 

08 

+ 

0. 

00 

0. 

05 

73 

I n  t  e  r  0  r  b  i  t  a  1    width 

0. 

1  1 

+ 

0. 

00 

0. 

33 

74 

Maxillary    length 

0. 

17 

+ 

0. 

01 

-0. 

35 

74 

Lower    jaw    length 

0. 

19 

+ 

0. 

00 

-0, 

01 

78 

Caudal   peduncle    dep 

th 

0. 

12 

+ 

0. 

00 

-0. 

61 

80 

Caudal   peduncle    len 

gth 

0. 

24 

+ 

0. 

01 

0. 

54 

81 

D2I 

0. 

05 

+ 

0. 

01 

1. 

69 

46 

All 

0. 

05 

+ 

0. 

01 

2. 

11 

56 

P2  I 

0. 

10 

+ 

0. 

01 

0. 

71 

75 

Pigmentation  variable  with  age  (see 
Ontogenetic  change ) ;  scale  pockets  mottled 
with  black;  fin  membranes  black;  opercular 
region  of  adults  black-slate  gray;  mouth 
primarily  light;  iris  black.  Specimens  fre- 
quently abraded,  underlying  tissue  pale 
yellow-rust  brown;  guanine  deposits  rare, 
if  present  occurring  on  opercular  complex, 
isthmus,  thorax,  or  abdomen;  silvered  forms 
generally  from  old  collections. 

Descriptions  based  on  104  specimens 
45.7-194.0  mm  SL. 

Onto<ienetic  chan2,e.  Maturation  in  E. 
pandionis  is  accompanied  by  changes  in 
pigmentation  and  dentition.  The  most 
striking  transformation  involves  caudal  pe- 
duncle markings.  Specimens  smaller  than  S5 
mm  SL  bear  a  thin,  black,  posterodorsally 
sloped  ring  circling  the  central  portion  of 
the  caudal  peduncle.  Melanophores  form- 
ing the  ring  are  deeply  embedded  in  pe- 
duncle musculature  and  are  not  easily 
abraded.   A  broader,  more  superficial  band 


of  pigment  circles  the  caudal  peduncle  at 
the  base  of  the  caudal  fin  (Fig.  IB).  As 
specimens  grow  beyond  85  mm,  the  rings 
become  fainter  and  begin  to  disappear. 
Fish  larger  than  110  mm  SL  may  com- 
pletely lack  peduncle  markings,  and  by 
125  mm  SL,  rings  are  absent  from  \irtually 
all  specimens.  Since  E.  pandionis  becomes 
sexually  mature  at  approximately  110  mm 
SL,  altered  markings  may  reflect  changes 
in  habit  or  behavior  associated  with  repro- 
ductive individuals. 

Gill  rakers  and  branchial  membranes  are 
converted  from  pale  yellow  to  black.  Spec- 
imens smaller  than  55  mm  SL  bear  scat- 
tered black  melanophores  on  gill  rakers 
but  lack  opercular  pigmentation.  B\-  60 
mm  SL  rakers  have  become  totalK'  dark, 
and  traces  of  melanin  ha\'e  appeared  on 
membranes  lining  the  opercle.  Pigment 
becomes  denser  with  growth  and  spreads 
ventrally.  By  100  mm  SL  the  opercle  is 
completely   lined   with   dark   tissue.    Since 


166       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Ei'iGONus  Systematics  •  Mayer       167 


opercular  bones  are  translucent,  the  process 
appears  outwardly  as  a  darkening  of  the 
opercle. 

Ontogenetic  changes  in  dentition  involve 
the  production  of  increasingly  complex 
tooth  patches.  Specimens  smaller  than  80 
mm  SL  bear  single  rows  of  teeth  on  the 
premaxillae  and  palatines.  Mandibular  teeth 
are  arranged  in  patterns  analogous  to  those 
found  on  the  premaxillae  or  in  double  rows 
that  taper  to  a  single  row  posteriorly. 
Vomerine  teeth  occur  in  1-2  chevron- 
shaped  clumps.  As  growth  takes  place, 
teeth  are  added  to  all  dentigerous  surfaces. 
Large  specimens  (>130  mm  SL)  have  as 
many  as  3-4  tooth  rows  on  palatines  and 
anterior  segments  of  dentaries  and  premax- 
illae. Vomerine  teeth  may  become  suf- 
ficiently numerous  to  cover  the  entire  face 
of  the  bone. 

Distribution.  E.  pandionis  is  amphi-At- 
lantic,  occurring  primarily  in  the  Caribbean, 
Gulf  of  Mexico,  and  Gulf  of  Guinea  (Fig. 
9).  The  species  has  been  taken  as  far 
north  as  New  Jersey  and  as  far  south  as 
French  Guiana  in  the  western  Atlantic. 
It  occurs  between  Portuguese  Guinea  and 
Angola  in  the  eastern  Atlantic.  Adults  are 
captured  exclusively  by  bottom  trawls  be- 
tween 210  and  600  meters.  American  forms 
are  most  numerous  from  300  to  500  meters, 
while  African  populations  are  most  abun- 
dant between  260  and  450  meters.  A  single 
pelagic  juvenile  (35.5  mm  SL,  MCZ  48839) 
was  taken  at  275  to  300  meters  in  the 
Caribbean. 

Geo<:,raphic  variation.  Statistical  analyses 
provide  conflicting  assessments  of  the 
similarity  of  African  and  American  popula- 
tions. Meristic  characters  reveal  little  vari- 
ability. Coefficients  of  difference  calculated 
for  standard  counts  are  always  less  than 
or  equal  to  0.49 — far  below  conventional 
levels  of  subspecies  recognition.  Mensural 
data,  on  the  other  hand,  suggest  there  are 
considerable  differences  between  the  pop- 
ulations. Of  thirteen  traits  analyzed,  seven 
separate  eastern  and  western  populations 
at  the  95%  level  of  confidence,  five  separ- 


ate them  at  the  98%  level,  and  two  separate 
them  at  the  99%  level  (Table  10). 

A  closer  examination  of  the  characters 
exhibiting  signilicant  differences  reveals 
that  regression  coefficients  of  American 
E.  pandionis  are  always  greater  than  those 
of  African  forms.  Since  regression  coef- 
ficients are  a  measure  of  relative  growth, 
observed  intraspecific  variation  may  reflect 
environmental  factors. 

Water  temperature  is  a  major  parameter 
determining  growth  rates  in  fishes.  If  other 
factors  are  conti'olled,  rates  of  growth  in- 
crease proportionally  with  temperature 
(Brown,  1957:  391).  With  this  in  mind,  it 
is  interesting  that  temperatures  are  gener- 
ally higher  and  superficial  warm-water 
layers  thicker  in  the  western  tropical  At- 
lantic (Ekman,  1953).  At  300  meters  Gulf 
of  Mexico  and  Caribbean  temperatures  vary 
from  10  to  18°  C  while  west  African  tem- 
peratures range  between  9  and  11°  C.  At 
500  meters  the  difference  is  slightly  less 
pronounced — 8-13°  C  as  opposed  to  6-8° 
C  (from  temperatin-e  profiles  in  Fuglister, 
1960;  Wiist,  1964;  and  Nowlin  and 
McLellan,  1967).  One  would  therefore 
expect  western  Atlantic  E.  pandionus  to 
grow  more  rapidly  and  exhibit  larger  re- 
gression coefficients  than  eastern  Atlantic 
forms.  In  view  of  these  findings,  the  two 
morphs  are  not  considered  to  represent 
separate  subspecies. 

Remarks.  See  E.  trewavasae:  Remarks 
for  discussion  of  E.  pandionis  .sensu  Lozano 
(1934),  Navarro  et  al.  (1943),  and  Maurin 
(1968). 

Specimens  of  doubtful  identity.  Five  spec- 
imens were  examined  that  resembled  E. 
pandionis  but  could  not,  with  certainty, 
be  placed  in  the  species.  Four  were  taken 
in  the  Atlantic,  the  fifth  in  the  Gulf  of 
Oman  (see  Mayer,  1972:  Appendix  II  for 
complete  data).  These  fishes  were  not 
considered  when  preparing  the  description 
of  E.  pandionis,  nor  were  they  used  in 
morphometric,  meristic,  or  distribution 
analyses. 

Tlie    Atlantic    specimens    include    three 


168       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  10.     Comparison  of  regression  coefficients  from  eastern  and  %vestern 
Atlantic  populations  of  E.  pandionis.    Data  evaluated  at  the  95%,  98%,  and 

99%  LE\rELS  OF  confidence.     DF   =:  DEGREES  OF  FREEDOM;   Eb   =   REGRESSION   COEFFI- 
CIENTS  OF   EASTERN   ATLANTIC    SPECIMENS;    SD    =    SIGNIFICANT    DIFFERENCE    BETWEEN 
TABULAR  AND  CALCULATED  VALUES  OF  t;   t=3   CALCULATED  VALUES   OF  t;   Wb   =   REGRES- 
SION COEFFICIENTS  OF  WTESTERN  ATLANTIC  SPECIMENS. 

Significance 
Wb  Eb         DF  t  S  S  S 


HL 

0. 

36 

0. 

37 

60 

0. 

97 

Body  depth 

0. 

29 

0. 

29 

71 

0, 

31 

Head  height 

0. 

22 

0. 

20 

63 

2. 

41 

SD 

SD 

Eye  diameter 

0. 

17 

0. 

15 

76 

2. 

14 

SD 

Snout  length 

0. 

09 

0. 

08 

69 

3. 

00 

SD 

SD 

SD 

Interorbital  width 

0. 

11 

0, 

11 

70 

2. 

13 

SD 

Maxillary  leng 

^th 

0. 

18 

0. 

16 

70 

3. 

35 

SD 

SD 

SD 

Lower  jaw  len 

gth 

0. 

20 

0. 

18 

74 

2. 

63 

SD 

SD 

SD 

Caudal  pedunc 
depth 

le 

0. 

13 

0. 

12 

76 

2. 

39 

SD 

SD 

Caudal  pedunc 
length 

le 

0. 

23 

0. 

24 

77 

1. 

16 

D2I 

0. 

05 

0. 

06 

32 

1 

83 

All 

0. 

05 

0. 

05 

52 

0 

91 

P2I 

0. 

10 

0. 

10 

71 

0 

83 

fishes  from  St.  Helena.    The  most  recently  for  only  the  Caribbean  form,  which  was 

collected    (UZM   P45148)    was   incorrectly  taken  at  relatively  shallow  depths.    Exact 

identified  as  E.  telescopus  by  Banchot  and  determination  of  the  variants'  status  must 

Blanc  (1961).  The  two  older  fonns  (BMNH  await  the  capture  of  additional  material. 

1868.3.11.14/15)     are    probably    the    fish  The     Indian     Ocean     form     (BMNH 

discussed    by    Giinther    (1868).     The    re-  1889.4.15.24)     is     distinguished     from     E. 

maining    specimen    (USNM    207703)    was  pandionis  by  its  shallow  body  (22.5%  SL), 

taken  in  the  Caribbean.  narrow  interorbital  region  (8.3%  SL),  den- 

The  four  Atlantic  individuals   are   basi-  tigerous  glossohyal,  numerous  weak  oper- 

cally   similar   to   E.   pandionis  but   exhibit  cular  spinelets,  and  elongate  gill  filaments, 

shallower  heads  ( 17.4-19.8%  SL),  narrower  The  last  trait  suggests  the  fish  may  have 

interorbital  regions    (8.6-9.4%   SL),   fewer  inhabited   an  oxygen  minimum  layer.    As 

pyloric  caeca  (8-9),  and  fewer  gill  rakers  with  the  Atlantic  variants,  additional  ma- 

( 25-27 ) .    In  these  respects  they  resemble  terial  must  be  collected  before  the  status  of 

E.  fragilis.   Little  is  known  about  the  habits  the  form  can  be  determined, 

of  the  variants;  station  data  are  available  Common  names.  None. 


Epigonus  Systematics  •  Mayer       169 


Z^A 


^ 
y 


y 


x» 


Figure  10.     Epigonus  fragilis,  HOLOTYPE,  89.1   mm  SL,  CM  3900/FMNH  55204  (from  Jordan  and  Jordan,  1922). 


Epigonus  fragilis  (Jordan  and 
Jordan,  1922) 
Figure  10 

Scepterias  fragilis  Jordan  and  Jordan,  1922:  45, 
plate  II,  fig.  2  (original  description;  Honolulu 
market;  holotype  examined,  CM  3900/FMNH 
55204). 

?Hynnodus  fragilis  Pietschmann,  1930:    13. 

Diagnosis.  E.  fragilis  most  closely  re- 
sembles E.  pandionis  but  may  be  dis- 
tinguished by  its  shallow  body  (18.8-21.1% 
SL)  and  short,  shallow  head  (length  31.7- 
34.0%  SL,  height  16.0-17.4%  SL).  Unlike 
£.  pandionis,  E.  fragilis  lacks  peduncle 
rings  on  specimens  smaller  than  100-120 
mm  SL. 

In  the  past  E.  fragilis  has  been  confused 
with  Hijnnodus  atherinoides,  a  junior  syn- 
onym of  E.  occidentalis.  E.  fragilis  may  be 
distinguished  on  the  basis  of  body  depth 
( see  above ) ,  pectoral  fin  counts  ( 16-17 ) , 
and  the  absence  of  a  pungent,  bony  oper- 
cular spine.  Weak  opercular  armor,  to- 
gether with  second  dorsal  fin  counts  of  1,10 
differentiate  E.  fragilis  from  E.  treivavasae, 
E.  pectinifer,  E.  rohustus,  E.  lenimen,  and 
E.  crassicaudus.  Gill  raker  counts  of  25-26 
separate  E.  fragilis  from  all  remaining  con- 
geners except  E.  telescopiis.  E.  fragilis  may 
be  distinguished  from  the  latter  by  the 
presence  of  7-8  pyloric  caeca. 

Description.  E.  fragilis  is  known  from 
only  five  specimens.  Of  these,  the  holotype 
is  of  little  descriptive  value.  The  specimen 
is  severely  dehydrated  and  has  become 
discolored,  brittle,  and  shrunken.    The  fol- 


lowing account  is  based  primarily  on  two 
recently  captured  specimens  of  E.  fragilis 
(LACM  32668-6  and  USNM  207704)  and 
two  forms  collected  by  D.  S.  Jordan  in  1921 
(SU  23246).  The  latter  are  mentioned  in 
the  original  description  of  E.  fragilis  but 
are  not  designated  as  types. 

All  meristic  and  mensural  data  are  pre- 
sented in  the  text.  Detailed  statistical 
analyses  were  not  undertaken  because  of 
small  sample  size. 

Body  elongate;  anterodorsal  profile  con- 
vex, rising  without  interruption  from  tip 
of  snout  to  first  dorsal  fin.  Body  depth 
18.8-21.1%  SL;  caudal  peduncle  length 
25.4-26.9%  SL. 

Head  short,  31.7-34.0%  SL;  head  height 
16.0-17.4%  SL;  snout  blunt,  7.2-7.9%  SL; 
angle  of  gape  moderate;  jaws  equal.  Max- 
illa reaching  %  eye  length;  posteriormost 
point  of  maxilla  at  ventral  edge  of  bone. 
Eye  round,  38.1—41.5%  HL;  anterodorsal 
rim  of  orbit  reaching  profile;  interorbital 
width  8.8-9.4%  SL. 

Dentition  variable  with  age.  Teeth  con- 
ical; premaxillary  teeth  in  irregular  double 
rows  anteriorly,  tapering  to  single  row 
posteriorly,  occupying  anterior  %-%  of 
bone.  Mandibular  dentition  more  promin- 
ent than  that  of  premaxilla;  teeth  recurved, 
occupying  from  %  to  entire  length  of 
dentary,  arranged  in  single  or  double  rows 
near  symphysis  and  single  row  posteriorly. 
Vomerine  teeth  recurved,  arranged  in  oval 
or  diamond-shaped  patch,  covering  entire 
face    of    bone    in    adults.     Palatine    teeth 


170       Bulletiti  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Figure  11.     Epigonus  occidentalis,  152.7  mm  SL,  MCZ  48840. 


medially  recurved,  arranged  in  single-triple 
rows  anteriorly,  tapering  to  single  row 
posteriorly;  tongue  edentulous. 

Opercular  spine  weak,  ventral  to  7-9 
small  serrae;  angle  of  preopercle  produced, 
rounded,  ornamented  with  striations  and 
weak  serrations;  subopercle  and  inter- 
opercle  unornamented.  Gill  rakers  25  (3), 
26  (1),  simple,  awl-like.  Pyloric  caeca 
7(1), 8(2). 

First  dorsal  fin  VII  (4),  VIII  (1); 
second  dorsal  fin  1,10  (5);  anal  fin  11,9 
(5);  pectoral  fin  16  (1),  17  (3);  DJ 
moderate  to  long,  5.9-8.9%  SL;  DJ  short, 
6.9%  SL;  PJ  long,  10.1-10.2%  SL;  All 
broken. 

Vertebrae  10  +  15  (4);  epipleural  ribs 
not  visible  on  radiographs;  pleural  ribs 
8  (4),  inserting  on  vertebrae  3-10.  Pored 
lateral  line  scales  49  ( 2 ) . 

Color  in  alcohol  yellow-brown;  fin  mem- 
branes dark;  iris  silver-black;  mouth  light; 
branchial  membranes  light,  darkening  with 
age. 

Distribution.  E.  fragilis  is  endemic  to 
the  Hawaiian  Islands  (Fig.  12).  The  spe- 
cies is  demersal  and  has  been  taken  between 
120  and  125  meters. 

Taxonomic  notes.  Six  years  after  E. 
fragilis  was  described,  Fowler  ( 1928 )  syn- 
onymized  the  species  with  a  second  Ha- 
waiian apogonid,  Hijnnodus  atherinoides 
Gilbert,  1905.  The  synonymy  achieved 
moderate  acceptance  and  appeared  in  sev- 
eral publications  (e.g.,  Matsubara,  1936; 
Tinker,  1944;  Gosline  and  Brock,  1960). 
Fowler's    conclusions    were    based    on    a 


33-mm  specimen  (BPBM  3914)  obtained 
by  the  Tanager  Expedition.  The  specimen 
is  in  extremely  poor  condition.  All  colora- 
tion has  been  lost,  most  of  the  muscle 
tissue  has  decomposed,  and  much  of  the 
skeleton  has  become  decalcified.  Although 
it  is  impossible  to  identify  the  fish  because 
of  its  condition,  the  following  traits  suggest 
it  is  neither  E.  fragilis  nor  H.  atherinoides: 
dorsal  fin  elements — VIII-1,8;  anal  fin  ele- 
ments— 11,6;  vertebrae — 11  +  14.  These 
data  differ  from  Fowler's  report  of  VI-I,8 
dorsal  elements,  no  anal  spines,  and  7  anal 
rays. 

As  was  discussed  in  the  diagnosis,  E. 
fragilis  is  distinct  from  H.  atherinoides. 
Fowler's  synonymy  appears  to  have  been 
based  on  inaccurate  data  taken  from  an 
incorrectly  identified  fish. 

Common  names.  None. 

Epigonus  occidentalis  Goode 
and  Bean, 1896 
Figure  11 

Epigonus  occidentalis  Goode  and  Bean,  1896:  233, 
plate  LXVI,  fig.  236  (original  description; 
Steamer  BLAKE,  off  Barbados,  237  fms.;  holo- 
type  examined,  MCZ  28032 ) . 

Hijnnodus  atherinoides  Gilbert,  1905:  618,  plate 
79  (original  description;  ALBATROSS  Sta. 
3867,  Pailolo  Channel,  Hawaii,  284-290  fms.; 
holotype  examined,  USNM  51601);  Jordan 
and  Jordan,  1922:  44;  Fowler  and  Bean,  1930: 
121. 

Hijnnodus  megalops  Smith  and  Radcliffe,  1912 
{in  Radcliffe,  1912):  445,  plate  38,  fig.  3 
(original  description;  ALBATROSS  Sta.  5388, 
12°51'30"N,  123°26T5"E,  between  Bnrias  and 
Luzon,  Philippines,  226  fms.;  holotype  ex- 
amined, USNM  70255). 


Ei'iGONus  Systematics  •  Mayer       171 


Table  11.     Epigonus    occidentalis   meristic    data.     X  =  mean;    SD 

STANDARP  nEVIMION;  n  =   NUMBER  OF  SPECIMENS. 


X 


Range 


SD 


Pectoral    fin    rays 
Gill    r  aker  s 
Lateral    line    scales 
Pyloric    caeca 


20. 

21 

19-2  1 

0.  59 

56 

24. 

68 

22-27 

1.  08 

60 

48. 

15 

46-51 

0.  97 

46 

9. 

27 

8-13 

1.  05 

45 

Table   12.     Epigonus  occidentalis  regression  data,    b   =   regression 

COEFFICIENT  ±  95%  CONFIDENCE  INTERVAL;  a  =  Y  INTERCEPT;   n  =   NUMBER 
OF  SPECIMENS.     AlL  REGRESSIONS  ON   SL. 


b 

a 

n 

HL 

0. 

34 

+ 

0. 

02 

0.  72 

48 

Body    depth 

0, 

19 

+ 

0. 

02 

-1.  72 

48 

Head    height 

0. 

15 

+ 

0. 

01 

0.  53 

49 

Eye    diameter 

0. 

16 

+ 

0. 

01 

0,  66 

49 

Snout    length 

0. 

08 

+ 

0. 

00 

0.  06 

49 

Interorbital    width 

0. 

08 

+ 

0, 

01 

0.  83 

39 

Maxillary    length 

0. 

13 

+ 

0, 

01 

0.  88 

51 

Lower    jaw    length 

0, 

15 

+ 

0. 

01 

1.  26 

51 

Caudal    peduncle    de 

pth 

0, 

10 

+ 

0. 

01 

-0.  90 

54 

Caudal   peduncle    len 

igth 

0. 

23 

+ 

0. 

01 

1.  50 

53 

D2  I 

0. 

05 

+ 

0. 

00 

1.  42 

34 

All 

0. 

05 

+ 

0. 

01 

2.  18 

42 

P2I 

0. 

09 

+ 

0. 

01 

0.  67 

47 

Diagnosis.  E.  occidentalis  is  distin- 
guished from  all  other  congeners  by  the 
combination  of  shallow  body  depth  (14.1- 
19.57t  SL),  reduced  gill  raker  counts  (22- 
27),  and  the  presence  of  a  pungent,  bony 
opercular  .spine.  It  is  frequently  confused 
with  E.  denticulatus. 

Description.    Meristic    values    presented 


in  Table   11;   regression  data  for  morpho- 
metric  traits  presented  in  Table  12. 

Body  elongate,  cigar-shaped;  anterodor- 
sal  profile  weakly  convex,  flattened,  rising 
gradually  from  tip  of  snout  to  interorbital 
region,  leveling  off  toward  occipital  region, 
and  rising  gradually  to  base  of  first  dorsal 
fin.     Body    depth    14.1-19.5%    SL,    body 


172       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


width  subequal  to  or  greater  than  body 
depth;  caudal  peduncle  narrow,  length 
22.4-28.1%  SL. 

Head  length  30.5-37.9%  SL;  head  height 
13.3-17.2%  SL;  angle  of  gape  moderate  to 
small;  lower  jaw  equalling  or  protruding 
slightly  beyond  upper  jaw.  Maxilla  reach- 
ing Vi-%  eye  length;  posterior  margin 
of  maxilla  moderate  to  narrow,  posterior- 
most  point  at  ventral  edge  of  bone.  Eye 
long,  oval,  40.6-52.3%  HL;  anterodorsal 
rim  of  orbit  reaching  or  projecting  into 
dorsal  profile;  interorbital  region  narrow, 
5.6-8.5%  SL. 

Teeth  conical;  premaxillary  and  man- 
dibular teeth  frequently  recurved,  arranged 
in  simple  single  row  or  single  row  widening 
to  double  or  triple  rows  near  symphysis; 
teeth  covering  %  to  entire  length  of  pre- 
maxilla  and  %  to  entire  length  of  dentary; 
vomerine  teeth  arranged  in  1-4  irregular 
rows;  palatines  rarely  edentulous,  teeth 
1-10,  arranged  in  single  row,  covering 
anterior  Vi-V^  of  bone;  tongue  edentulous. 

Opercular  spine  pungent,  bony,  ventral 
to  1-3  poorly  ossified  spinelets;  spine  sep- 
arated from  spinelets  by  shallow  indentation. 
Preopercular  angle  produced,  rounded  or 
pointed,  bearing  serrations  and  striations; 
subopercle  serrate,  occasionally  striate; 
interopercle  variable,  frequently  serrate. 
Gill  rakers  short,  awl-like. 

First  dorsal  fin  VII  (59);  second  dorsal 
fin  1,10  (59);  anal  fin  11,8  (1),  11,9  (59); 
DJ,  DJ,  All,  PJ  short,  equahing  1.1^.2%, 
4.8-7.8%,  4.8-9.2%,  and  8.0-11.3%  SL  re- 
spectively. 

Vertebrae  10  -I-  15  (35);  epipleural  ribs 
6  (19),  7  (5),  inserting  on  vertebrae  1-6 
or  1-7  respectively;  pleural  ribs  7  (31), 
8(1),  inserting  on  vertebrae  2-9  or  3-9. 

Color  in  alcohol  variable  with  preser- 
vation; skin  frequently  removed  by  trawl- 
ing; underlying  tissue  pale  yellow,  yellow- 
pink,  occasionally  marked  with  rust  brown; 
scale  pockets  and  fin  membranes  black; 
opercular  area  black-slate  gray,  occasion- 
ally tinged  with  silver;  lower  jaw,  bran- 
chiostegal    membranes,    and    thoracic    and 


abdominal  regions  occasionally  silvered; 
guanine  most  prevalent  on  specimens  from 
old  collections.  Mouth  color  variable  with 
age  (see  Ontogenetic  change);  iris  and 
branchial  region  dark. 

Description  based  on  62  specimens  58.2- 
178.9  mm  SL. 

Ontogenetic  change.  The  most  striking 
age-related  change  in  E.  occidentalis  is 
the  development  of  oral  pigmentation.  As 
in  E.  telescopus  and  E.  macro'ps,  immature 
forms  bear  pigmentless  or  slightly  pig- 
mented mouths,  while  adults  have  black- 
ened oral  membranes.  Pigmentation  first 
appears  in  specimens  80-110  mm  SL. 
Melanophores  develop  just  anterior  to  the 
pharynx  and  spread  rostrally,  covering  a 
third  of  the  roof  and  floor  of  the  mouth  and 
half  of  the  tongue  by  the  time  specimens 
reach  115-130  mm  SL.  By  150  mm  SL  the 
tongue  is  completely  black,  and  by  175- 
180  mm  the  entire  mouth  is  dark.  Branchial 
membranes  undergo  an  analogous  trans- 
formation before  specimens  reach  58  mm 
SL. 

A  faint  black  ring  circling  the  middle  of 
the  caudal  peduncle  was  observed  on 
three  small  E.  occidentalis  (<  65  mm  SL). 
Similar  markings  were  absent  from  larger 
individuals.  The  rings  are  reminiscent  of 
markings  observed  on  young  E.  macrops 
and  E.  pandionis  and  probably  represent 
a  juvenile  feature  that  is  lost  with  growth. 

Distribution.  E.  occidentalis  has  been 
taken  in  the  Caribbean,  Gulf  of  Mexico, 
and  western  tropical  Atlantic.  It  is  also 
known  from  the  Philippine  and  Hawaiian 
Islands  (Fig.  12).  The  species  is  caught 
by  bottom  trawls  between  360  and  735 
meters.  Adults  are  most  abundant  in  the 
Caribbean  from  500  to  550  meters. 

Geographic  variation.  E.  occidentalis,  as 
here  defined,  includes  two  nominal  species 
— Hijnnodus  atherinoides  Gilbert,  1905  and 
H.  megalops  Smith  and  Radcliffe,  1912. 
The  former  originally  represented  a  Ha- 
waiian endemic;  the  latter  represented  a 
Philippine  form.  In  1930  Fowler  and  Bean 
synonymized   the   Pacific   morphs.    In   the 


Epigonus  Systematics  •  Maxjer       173 


Figure  12.  Distribution  of  E.  fragilis  and  E.  occidentalis.  Map  A  shows  localities 
in  the  Caribbean  and  Gulf  of  Mexico.  Map  B  shows  localities  in  the  western 
Pacific.  J^  E.  fragilis,  individual  haul  of  demersal  adults;  %  E.  occidentalis,  indi- 
vidual haul  of  demersal  adults;  cross-hatching  indicates  areas  where  E.  occiden- 
talis are  frequently  taken. 


174       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


3  cm 


Figure  13.     Epigonus  denticulatus,  115.1  mm  SL,  UMML  12463. 


authors'  opinions,  characters  separating  the 
two  forms  were  "simply  minor  discrepancies 
of  portraiture  and  should  never  have  been 
credited  as  specific  distinctions   [p.  122] ." 

Although  descriptions  and  illustrations 
of  H.  atherinoides  and  H.  megalops  suggest 
a  link  with  E.  occidentalis,  detailed  com- 
parisons of  the  three  forms  were  never 
made.  To  a  large  extent  this  was  the  re- 
sult of  inadequate  sampling.  Until  the 
initiation  of  the  OREGON  cruises  in  1950, 
few  E.  occidentalis  were  available  for  study. 
Pacific  forms  are  still  poorly  represented; 
only  seven  specimens  have  been  collected. 
Reports  of  additional  material  by  Fowler 
(1928),  Matsubara  (1936),  Smith  ( 1949a,b, 
1961),  Kamohara  (1952),  and  Moreland 
( 1957)  are  based  on  misidentifications. 

Comparisons  of  E.  occidentalis  and  the 
H.  atherinoides-H .  megalops  complex  pro- 
vide no  evidence  to  support  their  status 
as  separate  species.  Analyses  of  head 
length,  body  depth,  head  height,  eye  di- 
ameter, snout  length,  interorbital  and  max- 
illary widths,  caudal  peduncle  length  and 
depth,  and  All  and  PJ  lengths  reveal  no 
significant  differences  between  the  pop- 
ulations at  either  the  95%,  98%,  or  99% 
levels  of  confidence.  Meristic  data  also 
show  considerable  overlap  for  most  char- 
acters; however,  the  coefficients  of  dif- 
ference for  pyloric  caeca  and  gill  raker 
counts  are  above  conventional  levels  of 
subspecies  recognition  (1.68  and  1.99,  re- 
spectively).     In     addition,     Atlantic     and 


Pacific   populations    may   be   distinguished 
by  minor  qualitative  characters  such  as: 

(1)  short,  rounded  preopercular  angles 
in  Atlantic  forms;  longer,  pointed 
angles   in   Pacific  specimens; 

(2)  fusion  of  uroneurals  1  and  2  in 
Atlantic  forms  (based  on  3  alizarin 
preparations ) ;  separate  occurrence 
in  Pacific  forms  (based  on  1  alizarin 
preparation ) . 

On  the  basis  of  the  above  information, 
Atlantic  and  Pacific  forms  are  placed  in 
the  same  species  but  considered  members 
of  separate  subspecies.  Formal  description 
of  the  subspecies  must  await  the  capture 
of  additional  Pacific  specimens. 

Remarks.  A  single  unripe  female  E. 
occidentalis  (USNM  197353,  172.1  mm  SL) 
was  found  carrying  small  egg  masses  in  the 
anterior  portion  of  its  mouth  (anterior  to 
the  tongue  and  vomer).  The  masses  con- 
tained 125  oval  eggs  0.40-0.55  mm  in 
diameter.  The  presence  of  eggs  in  the 
mouth  of  an  Epigonus  is  of  interest,  be- 
cause several  shallow-water  apogonids  ex- 
hibit oral  brooding.  No  such  activity  has 
ever  been  reported  for  deep-sea  forms. 

Although  it  is  difficult  to  say  with 
certainty,  the  E.  occidentalis  eggs  are  prob- 
ably not  incubating  clutches,  but  rather 
non-apogonid  ova  ingested  during  trawling. 
Unlike  the  egg  masses  of  typical  oral 
brooding  apogonids,  those  found  in  E.  oc- 
cidentalis are  broken,  disrupted,  and  con- 
tain very  few  eggs.  An  84.9-mm  specimen 
of  Cheilodipterus  affinis  was  reported  in- 


Epigonus  Systematics  •  Mayer       175 


Table  13.     Epigonus  denticulatus  meristic  data.    X  =  mean;  SD 

STANDARD    DEVIATION;    n    =    NUMBER    OF    SPECIMENS. 

X  Range  SD  n 


Pectoral    fin    rays 

19. 

09 

18-20 

0.  56 

54 

Gill    rakers 

30. 

98 

28-  34 

1.  10 

58 

Lateral   line    scales 

48. 

12 

46-  49 

0.  76 

43 

Pyloric    caeca 

11. 

83 

10-  14 

0.  85 

42 

ciibating  21,000  eggs  0.35-0.4  mm  in  di- 
ameter (Smith  et  al.,  1971:  8-9).  The  ova 
fully  occupied  the  oral  and  branchial 
chambers  and  extensively  distended  the 
head.  These  conditions  were  not  observed 
in  E.  occidentalis. 

It  is  possible  that  the  eggs  represent  the 
remnants  of  a  larger  mass  that  was  spit 
out  and  partially  reingested.  However, 
were  this  the  case,  one  might  expect  to 
find  eggs  in  the  stomach  (Sakomoto,  1930) 
or  gill  rakers.  No  eggs  were  found  in 
either  region. 

Finally,  Breder  and  Rosen  (1966)  state 
that  eggs  of  oral  brooding  apogonids  are 
lield  together  by  fibers  attaching  to  one 
pole.  The  eggs  of  E.  occidentalis  are 
loosely  embedded  in  an  open  matrix  of 
fibers.  Grape-like  egg  clusters  character- 
istic of  Apogon  semilineatus  (Ebina,  1931: 
20 )  were  not  observed. 

Common  names.  None. 

Epigonus  denticulatus  Dieuzeide,  1950 
Figure  13 

Pomatomus    telescoptis,    Vaillant    (in    part)     (not 

Risso,  1810),  1888:  376. 
Scepterias  lenimcn,  Whitley   (in  part)    (not  Whit- 
ley,    1935),     1935:     230;    Whitley     (in    part), 

1940:  420. 
Epigonus    atherinoides,    Matsubara    (not    Gilbert, 

1905),    1936:    120,  fig.    lA;   Smith,    1961:    378, 

fig.  3;  Kamohara,  1952:  37,  fig.  31. 
Hynnodus  atherinoides.  Smith  (not  Gilbert,  1905), 

1949a:  101;  Smith,  1949b:  210,  fig.  495A. 
Epigonus  denticulatus   Dieuzeide,    1950:    89,   figs. 

1-2     (original    description;    Algerian    Coast    at 


200-500  m;  holotype  not  examined);  Tortonese, 
1952:  72,  1  fig.;  Dieuzeide  et  al.,  1953:  216, 
2  figs.;  Tortonese  and  Queirolo,  1970:  33, 
fig.  6. 

Diagnosis.  E.  denticulatus  lacks  a  fully 
ossified  opercular  spine,  bearing  instead 
3-7  membranous  projections.  This  feature 
distinguishes  it  from  E.  occidentalis,  E. 
treivavasae,  E.  pectinifer,  E.  rohustus,  E. 
lenimen,  and  E.  crassicaudtis,  which  have 
pungent,  bony  opercular  spines.  E.  denti- 
culatus is  differentiated  from  E.  telescopus, 
E.  macrops,  and  E.  fragilis  by  the  presence 
of  10-14  pyloric  caeca  and  28-34  gill 
rakers.  It  differs  from  E.  oligolepis  by 
bearing  46-51  lateral  line  scales.  E.  denti- 
culatus closely  resembles  E.  pandionis  but 
may  be  distinguished  on  the  basis  of  the 
former's  shallow  body  (15.8-23.67r  SL), 
long  caudal  peduncle  (25.9-32.2%  SL), 
and  short  DJ  (2.4-3.7%  SL). 

Description.  Meristic  values  presented 
in  Table  13;  regression  data  for  morpho- 
metric  traits  presented  in  Table  14. 

Body  fusiform,  slightly  compressed;  an- 
terodorsal  profile  rising  gradually  above 
snout,  becoming  steeper  and  slightly  con- 
vex over  eyes,  thereafter  rising  gradually 
to  first  dorsal  fin;  body  moderate  to  shal- 
low, depth  15.8-23.6%  SL;  caudal  peduncle 
narrow,  length  25.9-32.2%o  SL. 

Head  moderate  to  short,  31.2-38.6%  SL; 
head  height  16.0-19.8%  SL;  snout  short, 
blunt;  angle  of  gap(>  moderate  to  large; 
lower  jaw  protruding  slightly  beyond  up- 
per jaw.    Maxilla  reaching  %-%  eye  length, 


176       Biilletifi  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  14.     Epigonus  denticulatus  regression  data,    b   =   regression 

COEFFICIENT  ±  95%  CONFIDENCE  INTERVAL;  a  =  Y  INTERCEPT;   11  =   NUMBER 
OF  SPECIMENS.    AlL  REGRESSIONS  ON  SL. 


HL 

0. 

32 

+ 

0. 

01 

2. 

88 

57 

Body    depth 

0. 

25 

+ 

0. 

01 

-4. 

09 

54 

Head    height 

0. 

16 

+ 

0. 

01 

0. 

86 

56 

Eye    diameter 

0. 

14 

+ 

0. 

01 

1. 

39 

58 

Snout    length 

0. 

07 

+ 

0. 

00 

0. 

33 

56 

Interorbital    width 

0. 

09 

+ 

0. 

00 

0. 

37 

55 

Maxillary    length 

0. 

14 

+ 

0. 

01 

1. 

37 

56 

Lower    jaw    length 

0. 

15 

+ 

0. 

01 

1. 

61 

57 

Caudal    peduncle    d 

epth 

0. 

11 

+ 

0. 

,  01 

-0. 

78 

57 

Caudal    peduncle    li 

ength 

0. 

28 

+ 

0. 

,  01 

0. 

61 

57 

D2I 

0. 

05 

+ 

0. 

,  01 

2. 

24 

36 

All 

0. 

,  06 

+ 

0, 

,  01 

1. 

,  43 

40 

P2I 

0. 

,  08 

a. 

0, 

,  01 

0. 

,  92 

41 

posteriormost  point  near  ventral  surface 
of  bone.  Eye  round  or  slightly  oval,  40.3- 
48.0%  HL;  anterodorsal  rim  of  orbit 
reaching  dorsal  profile,  projecting  into  pro- 
file in  smaller  specimens;  interorbital 
width  8.2-10.4%  SL. 

Teeth  small,  conical,  occasionally  re- 
curved; premaxilla  bearing  single  row  of 
teeth  along  anterior  Vs-%  (usually  %)  of 
bone.  Mandibular  teeth  arranged  along 
length  of  dentary  in  irregular  single  row, 
occasionally  double  near  symphysis;  larger 
specimens  with  3-4  rows  near  symphysis. 
Vomerine  teeth  variable,  arranged  in  1-4 
irregular  rows.  Palatine  dentition  occupy- 
ing length  of  bone,  arranged  in  simple 
single  row  or  double  row  tapering  to  single 
row  posteriorly;  large  specimens  bearing 
3-4  rows  of  teeth  anteriorly.  Tongue  gen- 
erally edentulous,  rarely  Ijearing  isolated 
tooth  patches  on  glossohyal  or  edges  of 
tongue. 

Opercle  lacking  bony  spine,  bearing  in- 


stead 3-7  (usually  5-6)  jagged,  mem- 
branous projections;  projections  often  ob- 
scured by  underlying  tissues.  Peropercular 
angle  produced,  broadly  rounded,  striations 
radiating  from  inner  edge,  angle  occasion- 
ally serrate;  subopercle  and  interopercle 
occasionally  serrate.  Gill  rakers  simple, 
awl-like. 

First  dorsal  fin  VII  (53);  second  dorsal 
fin  1,9  (1),  1,10  (56),  10  (1);  anal  fin  11,8 
(1),  11,9  (57).  DJ  moderate,  2.4-3.7% 
SL;  Dol,  All,  P,I  short,  5.2-8.0%,  6.0-8.2%, 
7.9-10.0%  SL  respectively. 

Vertebrae  10  +  15  (44);  epipleural 
ribs  6  ( 32 ) ,  7  ( 1 ) ,  inserting  on  vertebrae 
1-6  or  1-7  respectively;  pleural  ribs  8  ( 44 ) , 
inserting  on  vertebrae  3-10. 

Color  in  alcohol  variable  with  preserva- 
tion; skin  frequently  removed  by  trawling, 
underlying  tissue  pink-brown  or  yellow; 
scale  pockets  mottled  with  numerous 
brown-l)lack  melanophores,  dorsal  surfaces 
of  body  and  head  more  heavily  pigmented. 


Epigonus  Systematics  •  Mayer       177 


Guanine  deposits  frequently  occurring  on 
gill  cover,  ventral  surface  of  mandible, 
isthmus,  thoracic  region,  and  abdomen  to 
anus;  iris  black;  mouth  light;  branchial 
region  dark. 

Description  based  on  58  specimens  57.0- 
187.5  mm  SL. 

Ontogenetic  change.  Two  young  spec- 
imens of  E.  dcnfictihifus  (29.2  mm  SL, 
MCZ  48846,  and  49.7  mm  SL,  MCZ  48847) 
were  examined  in  the  course  of  this  in- 
vestigation. These  specimens  were  taken 
by  midwater  trawls  made  in  the  central 
North  Atlantic  and  Gulf  of  Mexico  and 
reveal  that  the  life  cycle  of  E.  denticulatus 
includes  a  pelagic  juvenile  stage. 

The  pelagic  young  resemble  adults  in 
most  respects.  For  example,  the  juveniles 
bear  diagnostic  gill  raker  counts  and 
opercular  ornamentation.  However,  slight 
changes  in  body  shape  are  associated  with 
growth.  The  29.2  mm  specimen  has  a  more 
shallow  body,  shorter  head,  narrower  inter- 
orbital  region,  and  smaller  eyes  than 
demersal  adults.  Similar  trends  are  present 
but  less  apparent  in  the  larger  juvenile. 

Juvenile  dentition  patterns  are  basically 
like  those  of  adults  but  involve  fewer  and 
relatively  larger  recurved  teeth.  Oral  and 
branchial  regions  are  light  in  young  speci- 
mens.   The  latter  areas  darken  with  age. 

Distribution.  E.  denticulatus  is  the  only 
cosmopolitan  species  in  the  genus  (Fig. 
14).  Specimens  have  been  taken  from  the 
southwest  coast  of  Japan,  the  Gulf  of 
Mexico,  and  the  Caribbean.  In  addition, 
the  species  occurs  continuously  from  the 
western  Mediterranean,  south  along  the 
western  coast  of  Africa  to  the  tip  of  the 
continent.  It  reappears  south  of  the  Great 
Australian  Bight  and  southeast  of  New 
Zealand. 

Adults  are  generally  taken  by  bottom 
trawls  between  300  and  600  meters,  al- 
though specimens  have  been  captured  from 
as  shallow  as  200  meters  and  as  deep  as  830 
meters.  Pelagic  juveniles  have  been  taken 
by  IKMT  between  130  to  145  meters  and 
350  to  425  meters. 


Geographic  variation.  E.  denticuhitus 
may  be  divided  into  North  Atlantic,  South- 
ern Hemisphere,  and  Japanese  populations. 
North  Atlantic  forms  include  material  from 
the  Mediterranean,  northeast  Atlantic, 
Caribbean,  and  Gulf  of  Mexico.  Southern 
Hemisphere  populations  contain  specimens 
from  the  southeast  Atlantic,  Australia,  and 
New  Zealand. 

Statistical  analyses  reveal  surprisingly 
little  divergence  between  North  Atlantic 
and  Southern  Hemisphere  specimens.  Co- 
efficients of  difference  for  standard  meristic 
characters  are  far  below  accepted  levels 
for  subspecies  recognition  (all  are  ^  0.53), 
and  regression  coefficients  for  mensural 
data  are  virtually  identical.  Only  maxil- 
lary lengths  differ  significantly  at  the  95% 
level  of  confidence.  It  is  clear  from  the 
data  that  North  Atlantic  and  Southern 
Hemisphere  E.  denticulatus  do  not  repre- 
sent separate  subspecies. 

Detailed  analyses  of  the  Japanese  pop- 
ulation could  not  be  undertaken  because 
of  inadequate  sampling.  Only  one  speci- 
men was  available  from  the  area.  On  the 
basis  of  this  fish,  the  Japanese  population 
appears  closely  allied  to  the  rest  of  the 
species.  With  the  exception  of  eye  di- 
ameter, standard  counts  and  measm-ements 
made  on  the  Japanese  morph  fall  within 
the  95%  and  99%  confidence  intervals  of 
remaining  E.  denticulatus.  Eye  diameter 
falls  outside  the  95%  confidence  interval 
but  within  the  99%  confidence  interval. 

The  similarity  of  E.  denficuhiius  pop- 
ulations, despite  the  wide  rangc>  of  the 
species,  suggests  ( 1 )  there  may  be  con- 
siderable gene  flow  between  populations, 
(2)  the  present  distribution  may  have  been 
achieved  only  recently,  or  (3)  evolution 
is  occurring  very  slowly.  Discovery  of  a 
pelagic  juvenile  in  the  mid-North  Atlantic 
gives  credence  to  the  first  hypothesis 
and  proN'ides  a  mechanism  for  the  dis- 
persal of  a  species  with  demersal  adults 
such  as  E.  denticulatus. 

Common  names.  "Castagnera  briina"  in 
Monaco   (Bini,  1968). 


178       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


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Epigonus  Systematics  •  Mayer       179 


Epigonus  oligolepis  sp.  nov. 
Figure  15 

llolotype:  One  specimen,  90.8  nun  SL,  taken 
from  the  Straits  of  Florida  bv  M/V  COMBAT, 
Sta.  436:  21  July  1957,  1319  to  1530  hrs.; 
24°13'N,  81°42'W;  300  fnis.,  10'  flat  trawl. 
USNM  207718. 

Parat>pes:  One  specimen,  126.7  mm  SL,  M/V 
OREGON,  Sta.  4731:  27  February  1964; 
27°35'N,  92°32'W;  250-300  fms.;  40'  flat 
trawl.  MCZ  48848. 

Three  specimens,  52.7-72.7  mm  SL,  Steamer 
ALBATROSS,  Sta.  2643:  9  April  1886; 
25°25'00"N,  79°55'15"W;  211  fms.  USNM 
109430. 

Three  specimens,  53.7-84.2  mm  SL,  M/V 
OREGON,  Sta.  5043:  26  September  1964; 
12°01'N,  6P53.5'W;  210-250  fms.;  40'  shrimp 
trawl.  USNM  207719. 

One  specimen  (cleared  and  stained),  62.0 
mm  SL,  locality  data  identical  with  those  of 
preceding  lot.  USNM  207720. 

One  specimen,  117.1  mm  SL,  M/V  OREGON, 
Sta.  3741:  26  August  1962;  29°10'N,  88°01.5' 
W;  300-340  fms.;  100'  flat  trawl.  USNM 
207721. 

Diagnosis.  E.  oligolepis  is  distinguished 
from  all  congeners  by  lateral  line  scale 
counts  of  33-36  and  the  presence  of  lingual 
and  endopterygoid  teeth. 

Description.  Meristic  values  presented  in 
Table  15;  regression  data  for  morphometric 
traits  presented  in  Table  16. 

Body  elongate,  moderately  compressed; 
anterodorsal  profile  rising  gradually  from 
tip  of  snout  to  interorbital  region,  rising 
more  steeply  and  becoming  slightly  convex 
to  occiput,  thereafter  rising  gradually  to 
base  of  first  dorsal  fin;  body  depth  19.8- 
24.5%  SL;  caudal  peduncle  length  23.9- 
27.2%  SL. 

Head  moderate  to  long,  34.4-43.0%  SL; 
head  height  16.6-18.8%  SL;  snout  pointed; 
angle  of  gape  moderate;  lower  jaw  pro- 
truding beyond  upper  jaw.  Maxilla  reach- 
ing %-%  eye  length;  posterior  margin 
of  maxilla  rounded,  posteriormost  point 
between  midline  and  ventral  margin  of 
bone.  Eye  round  to  slightly  oval,  40.1- 
43.77o  HL;  anterodorsal  rim  of  orbit  reach- 
ing or  projecting  into  dorsal  profile;  inter- 
orbital width  8.5-9.6%  SL. 


Teeth  small,  conical;  premaxilla  edent- 
ulous or  bearing  few  teeth  on  anterior  Vi- 
-f.  of  bone;  mandibular  teeth  arranged  in 
single  or  double  row  antericnly,  single  row 
posteriorly;  teeth  covering  anterior  half 
of  bone  and  occasionally  extending  along 
length  of  dentaiy.  Vomer  covered  with 
irregular  tooth  patches,  teeth  extending 
posteriorly  along  midline  of  palate;  pala- 
tine teeth  arranged  in  single  or  multiple 
rows  anteriorly,  single  row  posteriorly, 
covering  from  half  to  entire  length  of  bone; 
endopterygoid  dentigerous;  auxiliary  tooth 
patches  occurring  between  vomer,  pala- 
tines, and  endopterygoids;  tongue  den- 
tigerous, bearing  lateral  and  glossohyal 
tooth  patches  (Fig.  lA). 

Opercular  spine  weak,  poorly  ossified, 
ventral  to  2-6  membranous  spinelets;  spine 
and  spinelets  separated  by  moderate  gap; 
spinelets  occasionally  obscured  by  under- 
lying membranes.  Preopercular  angle  rec- 
tangular or  slightly  produced;  preopercle, 
subopercle  and  interopercle  unserrated. 
Gill  rakers  simple,  awl-like. 

First  dorsal  fin  VII  (10);  second  dorsal 
fin  1,10  (10);  anal  fin  11,8  (1),  11,9  (9). 
Fin  spines  moderate;  DJ  2.7-4.0%  SL; 
D,I  10.9-12.1%  SL;  All  10.3-12.2%  SL; 
PJ  11.0-13.6%  SL. 

Vertebrae  10  +  15  ( 10 ) ,  epipleural  ribs 
7  (4),  8  (1),  inserting  on  vertebrae  1-7 
or  1-8  respectively;  pleural  ribs  7  (10), 
inserting  on  vertebrae  3-9. 

Color  in  alcohol  variable  with  preserva- 
tion; specimens  frequently  abraded  reveal- 
ing underlying  pale  yellow  or  pink-purple 
tissue.  Recently  collected  specimens  bear 
scale  pockets  mottled  with  numerous 
melanophores;  dorsal  surfaces  of  head  and 
trunk  more  heavily  pigmented;  iris  black. 
Specimens  from  old  collections  devoid  of 
melanin,  bearing  silver  on  opercular  region, 
isthmus,  thoracic  region,  and  abdomen  to 
anus;  iris  silver.  Mouth  light,  dotted  with 
brown  or  black  melanophores;  l)ranchial 
region  light  in  small  specimens,  darkening 
with  age. 


180       Bulletin  Museum  of  Comparative  Zoology,  Vol  146,  No.  3 


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Epigonus  Systematics  •  Mayer       181 


Table  15.     Epigonus  oligolepis  meristic  data.   X  =  mean;  SD  =  stan- 
dard DEVIATION;  n  =  NUMBER  OF  SPECIMENS. 


X 


Range 


SD 


Pectoral    fin    rays  17.20  16-18  0.79  10 

Gill    rakers  30.50  29-31  0.71  10 

Lateral   line    scales  34.70  33-36  1.06  10 

Pyloric    caeca  8.83  8-10  0.75  6 


Table  16.     Epigonus  oligolepis  regression  data,  b  =  regression  coef- 
ficient ±  95%  confidence  interval;  a  =  Y  intercept;  n  =  number  of 
specimens.   All  regressions  on  SL. 


b 

a 

n 

HL 

0. 

36 

+ 

0. 

1 1 

0. 

88 

7 

Body    depth 

0. 

26 

+ 

0. 

02 

-2. 

45 

10 

Head    height 

0. 

21 

+ 

0. 

06 

-2. 

56 

5 

Eye    diameter 

0. 

15 

+ 

0. 

03 

0. 

48 

9 

Snout    length 

0. 

08 

+ 

0. 

03 

0. 

97 

5 

Interorbital    width 

0. 

10 

+ 

0. 

01 

-0. 

51 

9 

Maxillary    length 

0. 

18 

+ 

0, 

02 

-0. 

92 

6 

Lower    jaw    length 

0. 

17 

+ 

0. 

02 

1. 

45 

10 

Caudal    peduncle    d 

epth 

0. 

11 

+ 

0. 

02 

-1. 

20 

10 

Caudal    peduncle    L 

e  n  g  t  li 

0. 

26 

+ 

0. 

04 

0. 

58 

9 

D2I 

0. 

13 

+ 

0, 

01 

-1. 

07 

6 

All 

0. 

12 

+ 

0. 

02 

-0. 

33 

10 

P2I 

0. 

12 

+ 

0. 

03 

0. 

13 

9 

Description  based  on  10  specimens  53.7- 
126.7  mm  SL. 

Ontogenetic  change.  Two  juvenile  E. 
oligolepis  (32.0-32.2  mm  SL,  USNM  207722) 
were  taken  by  bottom  trawls  from 
the  Gulf  of  Mexico.  These  specimens 
exhibit  many  traits  characteristic  of  adult 
forms  but  differ  in  head  shape,  meristics. 


and  dentition.  Unlike  adults,  young  E. 
oligolepis  have  smaller  eyes  (38.2-39.4% 
HL)  and  wider  interorbital  regions  (10.4% 
SL).  Dorsal  fin  and  gill  raker  counts  are 
reduced  to  VI-I,10  and  26  respectively. 
Premaxillary,  mandibular,  and  lingual  tooth 
patterns  are  similar  to  those  of  mature 
individuals,   but   dentition   associated  with 


182       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Figure  16.     Distribution  of  E.  oligolepis.   ■  individual  haul  of  demersal  adults;  □  individual 
haul  of  demersal  juveniles. 


the  roof  of  the  mouth  is  strongly  reduced. 
Vomers  and  palatines  are  edentulous  or 
bear  1-4  teeth;  auxiliary  tooth  patches  have 
not  developed.  Endopterygoid  teeth  are 
present  but  few  in  number,  relatively  long, 
and  medially  recurved. 

Distribution.  E.  oligolepis  is  endemic  to 
the  Gulf  of  Mexico-Caribbean  region  (Fig. 
16).  Specimens  have  been  taken  by  bot- 
tom trawls  between  380  and  660  meters. 

Remarks.  The  type  specimens  of  E. 
oligolepis  exhibit  two  seemingly  disparate 
color  patterns.  One  lot,  taken  in  1886  by 
the  ALBATROSS,  is  devoid  of  melanin  but 
bears  extensive  guanine  deposits.  Remain- 
ing fish,  all  more  recently  collected,  bear 
no  silver  but  are  dotted  with  numerous 
melanophores.  These  differences  are  arti- 
facts of  preservation. 

Specimens  collected  by  early  workers 
were  generally  placed  directly  into  ethanol, 
while  material  obtained  today  is  fixed  in 
10  percent  formalin    (Hubbs   and   Lagler, 


1958:  16-17).  When  ethanol  is  used  as  a 
fixative,  it  leaches  out  melanins  but  does 
not  affect  guanine  deposits.  Specimens 
become  pale,  but  silver  pigment  is  retained. 
Formalin  has  the  opposite  effect;  it 
blackens  melanophores  but  destroys  gua- 
nine crystals.  The  appearance  of  preserved 
specimens  is  thus  dependent  on  fixative 
composition,  concentration,  and  immersion 
time.  An  alcohol-formalin  mixture  con- 
taining one  tablespoon  of  full  strength 
formalin  per  two  gallons  of  6.5-75  percent 
ethanol  might  be  used  instead  of  conven- 
tional fixatives  to  preserve  both  guanine 
and  melanin  deposits  (Myers,  personal 
communication ) . 

Etymology.  Oligolepis  (Greek),  few 
scales,  from  oligos,  few,  and  lepis,  scale; 
a  noun  in  apposition,  refers  to  the  reduced 
number  of  lateral  line  scales  characterizing 
the  species. 

Common  names.  None. 


Epigonvs  Systematics  •  Mayer       183 


Figure  17.     Epigonus  trewavasae,  98.6  mm  SL,  USNM  207723. 


Epigonus  trewavasae  Poll,  1954 
Figure  17 

Glossamia    pandionis,    Lozano     (not    Goode    and 

Bean,    1881),    1934:    89;    Navarro,    1942:    202; 

Navarro  et  al.,  1943:  136,  plate  XXII,  fig.  A. 
Epigonus     trewavasae    Poll,     1954:     91,     fig.     27 

(original    description;    NOORDENDE    III    Sta. 

52,    06°08'S,    11°30'E,    280-290    m;    holotype 

examined,  IRSN  209). 
Epigojitis    pandionis,     Maurin     (not     Goode     and 

Bean,  1881),  1968:  69,  fig.  36. 

Diagnosis.  E.  trewavasae  is  most  likely 
to  be  confused  with  E.  robustus,  E.  leni- 
men,  E.  crassicaudus,  and  E.  pectinifer.  It 
is  distinguished  from  the  first  three  species 
by  vertebral  counts  of  10  +  15  and  the 
presence  of  glossohyal  and  lateral  lingual 
teeth.  The  fourth  form,  E.  pectinifer,  bears 
only  glossohyal  teeth  or  a  totally  eden- 
tulous tongue.  E.  trewavasae  may  be 
further  differentiated  from  E.  pectinifer  on 
the  basis  of  the  former's  30-35  awl-like 
gill  rakers  and  long,  pungent  Dol  and  All 
(12.7-16.5%  SL,  13.8-16.8%  SL  respec- 
tively). E.  trewavasae  is  unlike  remaining 
congeners  because  it  bears  a  pungent,  bony 
opercular  spine,  second  dorsal  fin  counts 
of  1,9,  and  pectoral  fin  counts  of  16-18. 

Description.  Meristic  values  presented 
in  Table  17;  regression  data  for  morpho- 
mctric  traits  presented  in  Table  18. 

Body  elongate;  anterodorsal  profile  flat, 
rising  without  interruption  from  snout  to 
base  of  first  dorsal  fin;  body  moderate  to 
deep,  23.1-27.0%  SL;  caudal  peduncle 
length  24.3-27.5%  SL. 


Head  length  33.7-38.1%  SL;  head  height 
16.6-18.7%^'  SL;  snout  pointed;  angle  of 
gape  small  to  moderate;  lower  jaw  pro- 
truding beyond  upper  jaw,  bearing  two 
nubs  on  anterior  surface  of  mandible. 
Maxilla  reaching  slightly  less  than  %  eye 
length;  posterior  margin  of  maxilla  narrow, 
rounded,  or  bearing  posteriormost  point 
near  midline  of  bone;  short,  pungent  mus- 
tache-like process  projecting  from  postero- 
ventral  surface  of  maxillary  head.  Eye 
round,  slightly  oval  in  younger  specimens, 
41.1-49.1%f  HL;  anterodorsal  rim  of  orbit 
reaching  profile;  interorbital  width  8.8- 
10.8%  SL. 

Dentition  variable  with  age  (see  Onto- 
genetic change);  teeth  conical,  small,  fre- 
quently microscopic,  present  on  premaxiL 
lae,  mandibles,  and  vomer;  palatines 
occasionally  edentulous;  tongue  bearing 
lateral  and  glossohyal  tooth  patches. 

Opercular  spine  pungent,  bony,  sur- 
mounted by  2-3  horny  spinelets;  spine  and 
spinelets  separated  by  large  gap;  spinelets 
often  obscured  by  underlying  opercular 
membranes.  Preopercular  angle  narrowly 
produced,  unserrated  or  bearing  serrations 
on  angle  and  ventral  surface  of  bone;  in- 
teropercle  and  subopercle  unserrated  or 
weakly  serrated.  Gill  rakers  simple,  awl- 
like. 

First  dorsal  fin  VII  (14);  .second  dorsal 
fin  1,9  (13),  1,10  (1);  anal  fin  11,9  (14); 
DJ  moderate,  2.4-3.2%  SL;  DJ,  All,  PJ, 


184       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  17.     Epigonus  trewavasae  meristic  data.    X   =  mean;   SD 

STANDARD  DEVIATION;  n  =:  NUMBER  OF  SPECIMENS. 


X 


Range  SD 


Pectoral    fin    rays 
Gill    rakers 
Lateral   line   scales 
Pyloric    caeca 


n 


17.  54  16-  18  0.  66  13 

33.  15  30-  35  1.  46  13 

47.  69  47-  49  0.  75  13 

7.  00  6-     8  0.  60  12 


Table  18.     Epigonus  trewavasae  regression  data,   b  =  regression  co- 
efficient ±95%  confidence  interval;  a  ^  Y  intercept;  n  =:  number  of 
specimens.    All  regressions  on  SL. 


b 

a 

n 

HL 

0.  38 

+ 

0.  03 

-2.  03 

13 

Body    depth 

0.  29 

+ 

0.  02 

-4.  43 

12 

Head    height 

0.  19 

+ 

0.  01 

-1,  26 

12 

Eye    diameter 

0.  17 

+ 

0.  02 

-0,  49 

13 

Snout   length 

0.  07 

+ 

0.  02 

1.  19 

13 

Interorbital    width 

0.  09 

+ 

0,  01 

1.  41 

13 

Maxillary    length 

0.  15 

+ 

0.  02 

0.  69 

13 

Lower    jaw    length 

0.  16 

+ 

0.  01 

0.  39 

13 

Caudal    peduncle    de 

pth 

0.  13 

+ 

0.  01 

-1.  79 

1? 

Caudal    peduncle   len 

igth 

0,  26 

+ 

0.  02 

-0.  08 

13 

D2  I 

0,  15 

+ 

0.  03 

-0.  32 

12 

All 

0.  18 

+ 

0.  03 

-0.  65 

11 

P2I 

0,  14 

+ 

0.  01 

0.  36 

13 

long,  pungent,  12.7-16.5%,  13.(8-16.8%, 
13.8-16.27^  SL  respectively. 

Vertebrae  10  +  15  (12);  epipleural  ribs 
6  (9),  7  (2),  inserting  on  vertebrae  1-6 
or  1-7  respectively;  pleural  ribs  7  (8),  8 
(4),  inserting  on  vertebrae  3-9  or  3-10 
respectively. 

Color  variable  with  presei'vation;  speci- 
mens   abraded,    revealing   underlying   yel- 


low to  yellow-pink  tissue;  fin  membranes 
dark;  scale  pockets  covered  with  dense 
brown  or  black  melanophores;  dorsal  sur- 
face of  trunk  more  heavily  pigmented  than 
ventral;  opercles  brown,  black,  or  slate 
gray;  guanine  deposits  occurring  occasion- 
ally on  opercular  region  and  from  isthmus 
to  bases  of  paired  fins;  iris  black  with  sil- 
ver highlights;   mouth  light;  branchial  re- 


Epigonus  Systematics  •  Mayer       185 


JO'  SO" 


Figure  18.  Distributions  of  E.  trewavasae  and  E.  pectinifer.  Large  map  shows  localities  in  the  Atlantic;  insert 
shows  localities  off  Japan.  E.  trewavasae:  ^  individual  haul  of  adults;  Q  individual  haul  of  juveniles;  cross- 
hatching  indicates  areas  of  capture  cited  in  the  literature.  £.  pectinifer:  ■  individual  haul  of  adults;  □  individ- 
ual haul  of  juveniles;  A  report  from  the  literature. 


gion  light  in  smiill  specimens,  becoming 
l)lack  with  age. 

Description  based  on  13  specimens  70.9- 
153.9  mm  SL. 

Ontogenetic  change.  The  most  striking 
ontogenetic  changes  in  E.  trewavasae  are 
associated  with  the  development  of  adult 
tooth  patterns.  Large  specimens  bear  ir- 
regular double  or  triple  rows  of  premaxil- 
lary  and  mandibular  teeth  that  taper  to  a 
single  row  posteriorly.  Vomers  are  covered 
with  minute  conical  teeth,  while  palatines 
are  either  edentulous  or  bear  single  to 
double  rows  of  teeth. 

Dentition  patterns  are  simple  in  small 
specimens  but  become  more  complex  as 
teeth  are  added  during  growth.  A  29.8- 
mm  juvenile  lacks  both  premaxillary  and 
mandibular  teeth.  By  70-75  mm  SL  teeth 
are  present  in  single  rows  on  the  jaws,  and 
by  145  mm  SL  adult  tooth  patterns  pre- 
vail. As  premaxillary  tooth  patches  widen, 
they    extend    posteriorly    and     eventually 


cover  the  first  half  of  the  bone.  Analogous 
expansion  occurs  in  vomerine  tooth 
patches. 

Distribution.  E.  tretcavasae  is  known 
from  equatorial  west  Africa,  northwest 
Africa,  and  the  western  Mediterranean 
(Fig.  18).  It  has  been  taken  by  bottom 
trawls  between  200  and  600  meters. 

Geographic  variation.  Statistical  com- 
parisons of  African  and  Mediterranean  E. 
trewavasae  were  not  made  because  of 
small  sample  size.  As  additional  material 
is  collected,  the  following  intraspecific 
differences  should  be  examined: 

( 1 )  vomerine  and  palatine  teeth  more 
strongly  developed  in  Mediterra- 
nean forms; 

(2)  chin  nubs  more  strongly  developed 
in  African  forms; 

(3)  preopercular  serrations  more 
strongly  developed  in  Mediterra- 
nean forms. 

Although  the  significance  of  thc\se  features 


186       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


is  unknown,  they  suggest  that  African  and 
Mediterranean  forms  may  represent  sepa- 
rate subspecies. 

Taxonomic  notes.  Pomatomichthys  con- 
stonciae  GigHoh,  1880  may  be  a  synonym 
of  E.  trewavasae  Poll,  1954.  See  E.  tele- 
scopus:  Taxonomic  notes,  for  a  discussion 
of  this  possibility. 

Remarks.  Dieuzeide  (1950:  104-105) 
reported  that  specimens  designated  as 
Glossamia  pandionis  hy  hozano  (1934)  and 
NavaiTO  et  al.  (1943)  were  actually  mis- 
identified  E.  denticulatus.  This  is  incor- 
rect. Lozano's  report  is  based  on  a  single 
specimen  (131  mm  total  length)  taken 
from  the  Catillian  coast.  Among  the  char- 
acters cited  for  this  fish  are  dorsal  fin 
counts  of  VII-1,9,  pectoral  counts  of  16, 
and  an  All  subequal  to  the  eye  diameter 
(p.  89).  All  of  these  are  characters  diag- 
nostic of  E.  trewavasae.  E.  denticulatus 
bears  10  rays  in  the  second  dorsal  fin,  18- 
20  pectoral  rays,  and  an  All  equalling  half 
the  eye  diameter. 

Navarro  et  al.'s  specimens  also  appear 
to  be  E.  trewavasae.  Altliough  no  descrip- 
tion is  provided,  the  account  includes  a 
photograph  (plate  XXII,  fig.  A)  that  shows 
the  fish  have  deep  bodies,  pungent  oper- 
cular spines,  and  long  D2I,  All,  and  P2rs. 
All  of  these  features  are  characteristic  of 
E.  trewavasae. 

More  recently,  Maurin  (1968)  mistook 
E.  trewavasae  for  E.  pandionis.  Propor- 
tional measurements  of  body  depth,  head 
height.  All,  and  P-I  made  on  Maurin's 
figure  36  (p.  69)  fall  within  ranges  char- 
acteristic of  E.  treioavasae;  however,  pub- 
lished gill  raker  counts  of  28-30  (p.  70) 
are  lower  than  expected. 

Common  names.  None. 

Epigonus  pectin! fer  sp.  nov. 
Figure  19 

Ilolotype:  A  114.3-mm  SL  specimen  taken  from 
the  Caribbean  west  of  Grenada  by  M/V 
OREGON,  Sta.  5043:  26  September  1964, 
12°01'N,  61°53.5'W,  210-250  fms.,  40'  shrimp 
trawl.  USNM  207725. 


Paratypes:  One  specimen,  97.4  mm  SL,  16 
September  1964,  Suruga  Bay,  commercial  trawl. 
ABE  64-2085. 

One  specimen,  100.6  mm  SL,  14-31  October 
1964,  Suruga  Bay,  commercial  trawl.  ABE 
64-2245. 

One  specimen,  99.8  mm  SL,  14-31  October 
1964,  Suruga  Bay,  commercial  trawl.  ABE 
64-2248. 

Two  specimens,  95.2-117.1  mm  SL,  station 
data  identical  with  those  of  holotype.  MCZ 
48850. 

One  specimen  (cleared  and  stained),  108.1 
mm  SL,  station  data  identical  with  those  of 
holotype.  MCZ  48851. 

One  specimen,  94.8  mm  SL,  R/V  PILLS- 
BURY,  Sta.  P-582:  23  May  1967;  21°10'N, 
86°18'W;  250-155  fms.;  10'  otter  trawl.  UMML 
30378. 

One  specimen,  111.2  mm  SL,  M/V  OREGON, 
Sta.  4405:  27  September  1963;  11°53'N, 
69°28"W;  215  fms.;  40'  flat  trawl.  USNM 
207726. 

Ten  specimens,  101.8-120.6  mm  SL,  station 
data  identical  with  those  of  holotype.  USNM 
207727. 

Nine  specimens  81.5-118.9  mm  SL,  station 
data  identical  with  those  of  holotype.  USNM 
207728. 

Two  specimens  (cleared  and  stained),  94.8- 
98  mm  SL,  station  data  identical  with  those 
of  holotype.  USNM  207729. 

Epigonus  rohiistiis,  Matsubara  (not  Barnard, 
1927),  1936:  121,  fig.  IB;  Kamohara,  1952: 
37. 

Diagnosis.  E.  pectinifer  is  characterized 
by  comb-like  gill  rakers  on  the  lower  half 
of  the  first  gill  arch.  This  feature,  together 
with  glossohyal  dentition  (present  in  most 
specimens)  and  vertebral  counts  of  10  + 
15,  differentiate  E.  pectinifer  from  E.  ro- 
htistus,  E.  lenimen,  and  E.  crassicaudus. 
E.  pectinifer  most  closely  resembles  E.  tre- 
ioavasae but  is  distinguished  by  less  exten- 
sive lingual  dentition,  fewer  gill  rakers 
(26-30),  and  shorter  DJ  and  All  (11.2- 
12.7%  SL  and  11.9-14.0%  SL  respectively). 
E.  pectinifer  may  be  separated  from  re- 
maining congeners  by  its  pungent,  bony 
opercular  spine,  second  dorsal  fin  counts 
of  1,9,  and  pectoral  fin  counts  of  15-18. 

Description.  Meristic  values  presented 
in  Table  19;  regression  data  for  morpho- 
metric  traits  presented  in  Table  20. 


Epigonus  Systematics  •  Mayer       187 


CM 

o 

CM 


3 


E 
E 


UJ 
Q. 

>- 

!^ 

< 

Q. 


O 
V> 
Q. 

in 

C 
O 


o> 


188       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  19.     Epigonus  pectinifer  meristic  data.   X  =  mean;  SD  =  stan- 
dard DEVIATION;   n  =   NUMBER  OF  SPECIMENS. 


Range  SD 


Pectoral    fin    rays  16.03  15-18  0.57  29 

Gill    rakers  27.59  26-30  0.98  29 

Lateral   line    scales  48.14  47-49  0.58  29 

Pyloric    caeca  6,  10  5-     7  0.  41  29 


Table  20.     Epigonus  pectinifer  regression  data,  b  =  regression  coef- 
ficient ±  95%  confidence  interval;  a  =  Y  intercept;  n  =  number  of 
specimens.  All  regressions  on  SL. 


b 

a 

n 

HL 

0. 

35 

+ 

0. 

05 

-2. 

21 

27 

Body    depth 

0. 

28 

+ 

0. 

03 

-4. 

95 

28 

Head    height 

0. 

18 

+ 

0. 

03 

-2. 

83 

19 

Eye    diameter 

0, 

17 

+ 

0. 

03 

-3. 

18 

28 

Snout   length 

0. 

09 

+ 

0. 

03 

0. 

03 

26 

Interorbital    width 

0. 

10 

+ 

0. 

03 

-1, 

84 

28 

Maxillary    length 

0. 

17 

+ 

0. 

04 

-2. 

10 

26 

Lower   jaw   length 

0. 

15 

+ 

0. 

03 

0. 

55 

28 

Caudal  peduncle   de 

pth 

0. 

84 

+ 

0, 

03 

1. 

88 

28 

C  audal  peduncle    length 

0. 

27 

+ 

0. 

04 

-0. 

22 

28 

D2I 

0. 

11 

+ 

0. 

02 

0. 

62 

24 

All 

0. 

12 

+ 

0. 

02 

0. 

32 

24 

P2I 

0. 

11 

+ 

0. 

02 

1. 

30 

28 

Body  elongate;  anterodor.sal  profile  flat 
or  slighdy  convex,  rising  withont  interrup- 
tion from  snout  to  base  of  first  dorsal  fin; 
body  depth  21.1-24.6%  SL;  caudal  pedun- 
cle narrow,  length  2.5.1-28.7%  SL. 

Head  short  to  moderate,  31.3-35.7%  SL, 
shallow,  14.2-16.9%  SL;  snout  wide, 
pointed;  angle  of  gape  small;  lower  jaw 
proti-uding     slightly    beyond    upper    jaw; 


nubs  at  anterior  end  of  mandible  paired, 
barely  discernible,  or  absent.  Maxilla 
reacliing  %-y2  eye  length,  posterior  margin 
narrow,  rounded,  or  bearing  posteriormost 
point  near  midline  of  bone;  short,  pun- 
gent, mustache-like  process  projecting 
from  posteroventral  surface  of  maxillary 
head.  Eye  round  or  slightly  oval,  38.7- 
45.4%   HL;   anteiodorsal   rim  of  orbit   not 


Epigonus  Systematics  •  Maijer       189 


reaching  profile;  intcrorl)ital  widtli  7.7- 
9.4%  SL. 

Teeth  small,  conical;  premaxilla  edentii- 
lons  or  bearing  teeth  anteriorly;  when 
present,  teeth  1-15,  arranged  in  single 
row.  Mandibular  teeth  covering  all  or  part 
oi  dentary,  arranged  in  single  row.  Vomer- 
ine teeth  strong,  arranged  in  tightly  packed 
()\'al  patch.  Palatines  edentulous  or  bear- 
ing teeth  anteriorly;  when  present,  teeth 
1-6,  arranged  in  single  row;  tongue  with 
glossohyal  teeth,  rarely  edentulous. 

Opercular  spine  pungent,  bony,  ventral 
to  2-3  horny  spinelets;  spine  and  spinelets 
separated  by  large  gap;  spinelets  occasion- 
ally obscured  by  underlying  membranes. 
Preopercular  angle  narrowly  produced, 
serrated;  subopercle  and  interopercle  un- 
serrated  or  weakly  serrated.  Gill  rakers 
pectinate,  bearing  nub-like  projections 
proximally  along  mesial  surfaces  (Fig. 
IC);  pectinate  structure  variable  in  extent, 
most  prominent  on  ventral  portions  of  gill 
arch. 

First  dorsal  fin  VII  (28);  second  dorsal 
fin  1,9  (29);  anal  fin  11,9  (29);  D,I 
short,  1.6-2.8%  SL;  D,I  moderate,  11.2- 
12.7%  SL;  All,  PJ,  11.9-14.0%  SL. 

Vertebrae  10  +  15  (29);  epipleural  ribs 
6  (17),  7  (13),  inserting  on  vertebrae  1-6 
or  1-7  respectively;  pleural  ribs  8  (29), 
inserting  on  vertebrae  3-10. 

Color  in  alcohol  brown-black;  fin  mem- 
branes black;  scale  pockets  covered  with 
densely  packed  melanophores;  skin  often 
abraded,  revealing  underlying  yellow-pink 
tissue;  iris  black;  branchial  region  white  to 
dark  gray;  mouth  light. 

Description  based  on  30  specimens  81.5- 
120.6  mm  SL. 

OntO(!,enetic  change.  A  33.8-mm  E.  pec- 
tinifer  was  taken  by  bottom  trawl  in  the 
Gulf  of  Mexico  (USNM  207731).  The 
specimen  appears  similar  to  adults  and  pro- 
vides little  evidence  of  ontogenetic  change. 
The  major  difference  is  the  presence  of  six 
rather  than  seven  first  dorsal  fin  spines. 

Distribution.  E.  pectinifer  is  known 
from  the  Caribbean  Sea,  Gulf  of  Mexico, 


and  eastern  coast  of  Japan  (Fig.  18). 
Specimens  were  taken  between  280  and 
550  meters. 

GeograpJiic  variation.  Definitive  com- 
parisons of  Japanese  and  American  E.  pec- 
tinifer were  not  undertaken,  because  only 
three  oriental  specimens  were  available  for 
study.  The  latter  forms  were,  however,  in- 
dividually compared  with  Amcnican  fish. 
The  analyses  revealed  virtually  no  differ- 
ences between  the  populations  aside  from 
a  slight  tendency  toward  broader  caudal 
peduncles  and  shorter  maxillae  and  man- 
dibles by  the  Japanese  specimens. 

Remarks.  A  teratological  specimen  of 
E.  pectinifer  was  taken  from  the  Yucatan 
Channel  (109  mm  SL,  UMML  30379).  The 
fish  was  captured  at  depths  characteristic 
of  E.  pectinifer  and  bears  diagnostic  traits 
such  as  27  gill  rakers  ( many  are  pectinate ) , 
VII  +  I  dorsal  fin  spines,  16  pectoral  fin 
rays,  and  10  +  15  vertebrae.  The  tongue 
is  edentulous.  Unlike  the  condition  in 
typical  forms,  opercles  are  not  fully  ossified 
and  lack  spines  and  spinelets.  Similarly, 
the  lateral  line  is  incomplete  on  the  right 
side  and  bears  only  43  pored  scales  on  the 
left.  Other  differences  include  enlarged 
teeth  and  chin  nubs,  10  rather  than  9  dor- 
sal rays,  and  8  rather  than  5-7  pyloric 
caeca. 

The  aberrant  specimen  was  not  consid- 
ered in  preparing  the  species  description. 

Etymology.  Pectinifer  (Latin),  comb- 
bearer,  from  pecten,  comb,  and  ferare,  to 
bear;  a  noun  in  apposition,  refers  to  the 
comb-like  gill  rakers  characterizing  this 
species. 

Common  names.  None. 

Epigonus  robustus  (Barnard,  1927) 
Figure  20 

Epigonus  macrops  Gilchrist  and  von  Bonde,  1924: 
14,  plate  I,  fig.  3  (oiij^inal  description;  S..S. 
PICKLE  Sta.  344,  30°12'00"S,  14°25'()()"E, 
510  fms.;  Sta.  347,  31°58'00"S,  16°00'00"E, 
670  fms.;  syntype  examined,  RUSI  669;  name 
suppressed,  junior  homonym  of  Oxi/odot}  macrops 
Brauer,  1906);  Barnard,  1927:'  523;  Smith, 
1961:  377,  fig.  2. 


190       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Figure  20.     Epigonus  robustus,  154.6  mm  SL,  LACM  11449-7. 


Parahynnodiis  robustus  Barnard,  1927:  525,  plate 
XXII,  fig.  4  (original  description;  off  Cape 
Point,  460  fms.;  holotype  in  poor  condition,  not 
examined,  SAM  13080). 

Hymwdus  robustus  Smith,  1949b:  210,  fig.  495. 

Diagnosis.  E.  robustus  sti-ongly  resem- 
bles E.  pectinifer,  E.  trewavasae,  and  E. 
lenimen.  It  may  be  distinguished  from 
the  former  two  species  by  the  absence  of 
hngual  teeth.  In  addition,  mihke  E.  pec- 
tinifer, it  has  awl-Hke  gill  rakers.  E.  ro- 
hu^us  differs  from  E.  lenimen  by  having 
a  narrow  interorbital  region  (6.5-8.2% 
SL),  short  DJ  (10.0-12.6%  SL)  and  short 
All  (9.2-13.3%  SL).  E.  robustus  may  be 
distinguished  from  E.  crassicaudus  by  the 
former's  short  head  (28.0-34.0%  SL)  and 
shallow  body  (20.3-24.6%  SL).  It  differs 
from  remaining  congeners  by  bearing  a 
pungent,  bony  opercular  spine,  vertebral 
count  of  11  +  14,  and  nine  rays  in  the 
second  dorsal  fin. 

Description.  Meristic  values  presented 
in  Table  21;  regression  data  for  morpho- 
metric  traits  presented  in  Table  22. 

Body  elongate,  moderately  compressed; 
anterodorsal  profile  weakly  convex,  rising 
without  interruption  from  tip  of  snout  to 
base  of  first  dorsal  fin;  body  depth  20.3- 
24.6%  SL;  caudal  peduncle  moderate  to 
long,  25.3-30.7%  SL. 

Head  short,  shallow,  length  28.0-34.0% 
SL,  height  14.8-16.3%  SL;  snout  short, 
pointed;  angle  of  gape  moderate  to  large; 
lower  jaw  protruding  beyond  upper  jaw, 
bearing  two  nubs  of  variable  prominence 


on  anterior  surface  of  mandible.  Maxilla 
reaching  ¥3-^/4  eye  length;  posterior  margin 
of  maxilla  narrow,  rounded  or  bearing 
posteriormost  point  near  midline  of  bone; 
small,  weak  mustache-like  process  project- 
ing from  j)osteroventral  surface  of  maxil- 
lary head.  Eye  round  to  oval,  small,  37.4- 
42.4%  HL;  anterodorsal  rim  of  orbit  not 
reaching  dorsal  profile;  interorbital  region 
narrow,  6.5-8.2%  SL. 

Teeth  small,  conical;  premaxilla  edentu- 
lous or  bearing  single  row  of  teeth  on  an- 
terior half  of  bone;  mandibular  dentition 
covering  all  or  part  of  dentary,  arranged 
in  double  row  anteriorly,  tapering  to  single 
row  posteriorly;  vomer  bearing  1-6  irregu- 
lar rows  of  teeth;  palatines  edentulous  or 
bearing  teeth  on  anterior  half  of  bone; 
tongue  edentulous. 

Opercular  spine  pungent,  bony,  ventral 
to  2-3  membranous  or  horny  spinelets; 
spine  separated  from  spinelets  by  wide 
gap;  spinelets  often  obscured  by  underly- 
ing membranes.  Preopercular  angle  not 
produced,  serrations  on  posterior  and/ or 
ventral  surfaces  of  bone  rarely  absent;  sub- 
opercle  and  interopercle  serrated.  Gill 
rakers  simple,  awl-like. 

First  dorsal  fin  VI  (1),  VII  (27),  VIII 
(1);  second  dorsiil  fin  I,  9  (28),  II,  8  (1); 
anal  fin  II,  9  (29).  DJ  short,  1.4-2.5% 
SL;  DJ,  All,  PJ  moderate  to  long,  10.0- 
12.6%,  9.2-13.3%,  11.7-15.3%  SL  respec- 
tively. 

Vertebrae  11  +  14  (29);  epipleural  ribs 


Epigonus  Systematics  •  Mayer       191 


Table  21.     Epigonus  robustus  meristic  data.    X  =  mean;  SD  =  stan- 
dard DEVIATION;  n  =  NUMBER  OF  SPECrMENS. 

X  Range  SD  n 

Pectoral    fin    rays  16.79        16-18         0.55  29 

Gill    rakers  31.68       30-33         0.93  29 

Lateral    line    scales         48.76       47-50        0.91  29 

Pyloric    caeca  6.36  5-     8         0.78  28 


Table  22.     Epigonus  robustus  regression  data,   b  =  regression  coef- 
ficient ±  95%  confidence  interval;  a  =  Y  intercept;  n  =  number  of 
specimens.  All  regressions  on  SL. 


b 

a 

n 

HL 

0. 

28 

+ 

0. 

02 

5, 

35 

28 

Body    depth 

0. 

28 

+ 

0. 

02 

-7. 

80 

28 

Head    height 

0, 

17 

+ 

0. 

02 

-  1. 

65 

20 

Eye    diameter 

0. 

11 

+ 

0. 

01 

44 

28 

Snout    length 

0. 

06 

+ 

0. 

02 

80 

23 

Interorbital    widtl: 

0. 

09 

+ 

0. 

01 

-  1. 

97 

28 

Maxillary    length 

0. 

12 

+ 

0. 

01 

14 

24 

Lower    jaw    length 

0, 

14 

+ 

0. 

01 

27 

27 

Caudal    peduncle    dep 

ith 

0. 

13 

+ 

0. 

01 

-  1. 

69 

28 

Caudal    peduncle    len 

gth 

0. 

25 

+ 

0. 

02 

5. 

32 

28 

D2I 

0. 

08 

+ 

o„ 

02 

5. 

49 

2  1 

All 

0. 

07 

+ 

0, 

02 

8„ 

18 

15 

P2I 

0. 

09 

+ 

0. 

02 

6. 

74 

24 

6   (2),  7   (8),  inserting  on  vertebrae   1-6  brown   or   black   niclanopliores;    brancliial 

or  1-7  respectively;  pleural  ribs  9  (29),  in-  region  black.    Body  very  oily;  body  cavity 

serting   on  vertebrae   3-11.  filled  with  rust  brown  fat  globules;  viscera 

Color    variable    with    preservation,    pale  and  swinibladder  often  completely  envel- 

yellow  to   rust  brown;   scale  pockets   out-  oped  in  fat. 

lined   by    small    black    or    brown    melano-  Description     based     on     29     .specimens 

phores;     opercular     region     tinged     with  121.1-198.0  mm  SL. 

black;  iris  black;  mouth  light,  mottled  with  Distribution.    Most  specimens  of  E.   ro- 


192       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


EriGONus  Systematics  •  Mayer       193 


Figure  22.     Epigonus  lenimen,  139.0  mm  SL,  UZM  P45165. 


biistus  liave  been  taken  by  bottom  trawLs 
between  800  and  1225  meters  off  south- 
eastern South  Ameriea,  South  Africa,  and 
AustraHa  (sec  Fig.  21).  One  specimen 
(ISH  430/71)  was  taken  by  a  deep  pelagic 
trawl. 

Geographic  variation.  No  investigation 
was  undertaken  because  insufficient  ma- 
terial was  available  from  South  Africa  and 
Australia. 

Taxonomic  notes.  Epigonus  macrops 
Gilchrist  and  von  Bonde,  1924  was  des- 
cribed from  two  syntypes;  the  larger  was 
19(S  mm  (SL?).  These  specimens,  together 
with  manv  others  collected  bv  the  Fish- 
cries  and  Marine  Biological  Survey,  were 
lost  while  being  transferred  to  the  South 
African  Museum.  A  portion  of  the  ma- 
terial was  subsequently  rediscovered  at 
Rhodes  University,  Grahamstown.  From 
the  contents  Smith  (1961:  378)  described 
a  specimen  that  he  believed  to  be  "Gil- 
christ and  \'on  Bonde's  type  of  macrops 
from  600  fathoms  off  St.  Helena  Bay."  This 
fish  was  re-examined  during  the  present 
study. 

Smith's  specimen  measures  162.2  mm  SL 
and  thus  cannot  be  the  larger  syntype; 
however,  it  conforms  to  the  descriptions 
and  proportions  supplied  by  Gilchrist  and 
von  Bonde  and  probably  represents  the 
smaller  type  for  which  no  length  was  pub- 
lished. 

An  unusual  aspect  of  the  syntypes  is  that 


the  locations  at  which  they  were  captured 
will  never  be  precisely  known.  The  speci- 
mens were  taken  at  different  stations.  Al- 
though these  are  recorded  in  both  the  orig- 
inal description  of  E.  macrops  and  in  the 
1921  report  of  the  Fisheries  and  Marine 
Biological  Survey  (Gilchrist,  1922),  neither 
account  specifies  which  data  are  associated 
with  which  syntype. 
Common  names.   None. 

Epigonus  lenimen  (Whitley,  1935) 
Figure  22 

Scepterias  lenimen  WHiitley  (in  part),  1935:  230 
(original  description;  Great  Australian  Biglit: 
south  from  Eucla,  350—450  fnis.;  holot\-pe 
examined,  AM  E3368);  Whitley,  1940:  420, 
fig.  33;  Wliitley  (in  part),  1968:  56. 

Epigonus  lenimen  Scott,  1962:   191,  1  fig. 

Diagnosis.  E.  lenitnen  is  distinguished 
from  E.  robustus  and  E.  crassicaudus  by 
its  broad  interorbital  region  (8.7-10.2% 
SL),  long  DJ  (14.9-18.7%  SL),  and  large 
eyes  (40.0-51.1%  HL).  It  is  further  dif- 
ferentiated from  E.  crassicaudus  by  shorter 
head  lengths  (32.7-36.67^  SL)  and  .shal- 
lower head  heights  (16.2-18.8%  SL).  E. 
lenimen  lacks  lingual  teeth  but  has  11  +  14 
vertebrae  and  thus  may  be  distinguished 
from  E.  trewavasae  and  E.  pectinifer.  Un- 
like remaining  congeners,  E.  lenimen  bears 
a  pungent,  bony  opercular  spine,  nine 
second  dorsal  fin  rays,  and  16-18  pectoral 
fin  rays. 


194       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  23.  Epigonus  lenimen  meristic  data.  X  =  mean;  SD  =  standard 
deviation;  n  =  nxtmber  of  specimens. 


X 


Range 


SD 


Pectoral    fin    rays  16.96  16-18  0.58  28 

Gill    rakers  30.29  28-34  1.27  24 

Lateral   line    scales  48.12  47-50  0.91  26 

Pyloric    caeca  7.33  7-     9  0.56  24 


Table  24.     Epigonus  lenimen  regression  data,   b  =  regression  coeffi- 
cient ±  95%  confidence  interval;  a  =  Y  intercept;  n  =  nltmber  of 
specimens.   All  regressions  on  SL. 


b 

a 

n 

HL 

0. 

35 

+ 

0, 

01 

-0. 

12 

29 

Body    depth 

0. 

28 

+ 

0. 

02 

-3. 

32 

27 

Head    height 

0, 

19 

+ 

0. 

01 

-1. 

08 

27 

Eye    diameter 

0. 

18 

+ 

0. 

01 

-1, 

60 

27 

Snout   length 

0. 

08 

+ 

0. 

01 

0. 

31 

27 

Interorbital    width 

0. 

10 

+ 

0, 

01 

-0. 

59 

26 

Maxillary    length 

0. 

16 

+ 

0. 

01 

0. 

08 

27 

Lower    jaw    length 

0. 

16 

+ 

0. 

01 

0. 

55 

28 

Caudal    peduncle    dep 

th 

0. 

11 

+ 

0. 

01 

-0. 

14 

28 

Caudal    peduncle    len 

gth 

0. 

24 

+ 

0. 

02 

2. 

26 

26 

D2I 

0. 

17 

+ 

0. 

02 

-0. 

79 

18 

All 

0. 

21 

+ 

0. 

02 

-2. 

36 

22 

P2I 

0. 

19 

+ 

0. 

01 

-2, 

13 

28 

Description.  MerLstic  values  presented  in 
Table  23;  regression  data  for  morphometric 
traits  presented  in  Table  24. 

Body  elongate;  anterodorsal  profile  flat 
or  weakly  concave,  rising  without  inter- 
ruption to  first  dorsal  fin,  more  steeply 
inclined  behind  occiput  in  large  specimens; 
body   moderate    to   deep,    21.5-27.5%    SL; 


caudal  peduncle  moderate  to  long,  23.6- 
29.3%  SL.  Head  length  32.7-36.6%  SL; 
head  height  16.2-18.8%  SL;  snout  moder- 
ately pointed;  angle  of  gape  moderate, 
variable  with  age;  lower  jaw  protruding 
slightly  or  not  at  all;  no  prominent  nubs  on 
anterior  surface  of  mandible.  Maxilla 
reaching  Vs-V2  eye  length;  posterior  margin 


Epigonus  Systematics  •  Mayer       195 


Table  25.     Comparison  of  E.  LENiMEy  paratypes  with  specimens  of  E.  lknimen  and  E. 

DENTICULATVS.     PaRATYPE   MERISTICS   REPORTED   AS   VALUE,    FOLLOWED   IN   PARENTHESES    HY    NUM- 
BER OF  SPECLMENS  EXHIBITING  THAT  VALUE.     RATIOS  ARE  EXPRESSED  AS   PERCENTAGES. 

E  .     1  e  n  i  m  e  n 
E_.    1  e  n  i  m  e  n  paratypes  E^.    denticulatus 

Dorsal    fin    rays  8-9  9(1),  10(11)  10 

Pectoral    fin    rays  16—18  19(6),  20(6)  18—20 

Vertebrae  11+14  10+15  10+15 

10(1),  11(3)  , „       ,  , 

Pyloric    caeca  7  —  9  12(7)  — 

BH/SL  21.5—27.5  18.4  —  21.7  15.8—23.6 

D2  I/SL  14.9—18.7  6.0—7.6  5.3—8.0 

AII/SL  13.0  —  20.8  6.2—7.1  6.0—8.2 

P2   I/SL  12.5  —  18.7  8.5—9.9  7.9—10.0 


of  maxilla  narrow,  rounded,  or  bearing  First  dorsal  fin  VII  (29);  second  dorsal 
posteriormost  point  near  midline  of  bone;  fin  1,8  (1),  1,9  (28);  anal  fin  11,8  (2),  11,9 
weak  mustache-like  process  projecting  from  (26);  DJ  moderate,  2.0-4. 17^  SL;  DJ, 
posteroventral  surface  of  maxillary  head,  All  long,  14.9-18.7%,  13.0-20.8%  SL  re- 
process occasionally  absent.  Eye  large,  spectively;  PJ  moderate  to  long,  12.3- 
oval,  40.0-51.1%  HL;   anterodorsal  rim  of  18.7%  SL. 

orbit   reaching    dorsal   profile;    interorbital  Vertebrae  11  +  14  (29);  epipleural  ribs 

width  8.7-10.2%  SL.  6  (6),  7  (12),  8  (2),  inserting  on  vertebrae 

Teeth  small,  conical;  premaxilla  eden-  1-6,  1-7,  or  1-8  respectively;  pleural  ribs 
tulous  or  bearing  single  row  of  teeth  oc-  9  (28),  inserting  on  vertebrae  3-11. 
cupying  anterior  half  of  bone.  Mandible  Color  in  alcohol  variable;  skin  often 
edentulous  or  bearing  single  row  of  teeth  abraded,  revealing  underlying  pale  pink- 
occupying  up  to  %  of  dentary;  tooth  row  yellow  tissue;  fin  membranes  and  scale 
occasionally  double  near  symphysis.  Vo-  pockets  mottled  with  numerous  black 
mer  edentulous  or  bearing  up  to  seventeen  melanophores;  head,  opercular  region,  and 
teeth  arranged  in  diamond-shaped  patch  fin  bases  deep  rust  brown.  Guanine  de- 
or  in  1-3  irregular  rows;  palatines  edentvi-  posits  variable,  occurring  on  ventral  por- 
lous  or  bearing  1-2  teeth  anteriorly;  tongue  tions  of  opercular  region,  isthmus,  pectoral 
edentulous.  and  pelvic  fin  bases,  and  abdomen  to  anus; 

Opercular  spine  pungent,  bony,  ventral  silver  chromatophores  on  dorsal,  anal,  pee- 
to  1-5  (usually  2)  membranous  or  horny  toral,  or  pelvic  fin  rays;  iris  black  with 
spinelets;  spine  and  spinelets  separated  by  silver  highlights;  moutli  light,  dotted  with 
wide  gap;  spinelets  frequently  obscured  by  melanophores;  branchial  region  light  in 
underlying  membranes.  Preopercular  angle  small  specimcMis,  blackening  with  age. 
narrowly  produced,  occasionally  serrated;  Description  based  on  32  specimens  40.0- 
subopercle  and  interopercle  unserrated  or  147.8  mm  SL. 

weakly  serrated.    Gill  rakers  simple,  awl-  Distrihution.    E.  lenimen  is  known  from 

like.  three    localities    (Fig.    21).     The    liolotype 


196       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


0    I    2    3  cm 


Figure  23.     Epigonus  crassicaudus,  259.0  mm  SL,  MCZ  48855. 


was  taken  south  of  Austi-alia  between  622 
and  823  meters.  Remaining  specimens 
were  taken  off  New  Zealand  between  530 
and  660  meters. 

Taxonomic  notes.  Although  G.  P.  Whit- 
ley was  the  first  to  describe  E.  lenimen 
(1935),  inaccuracies  in  his  papers  have 
produced  several  problems.  The  most 
serious  involve  the  type  series  and  type 
locality  of  the  species. 

The  original  description  of  E.  lenimen 
designates  a  holotype  and  nine  paratypes. 
A  figure  of  the  new  form  was  not  included 
but  was  published  in  a  subsequent  paper 
(Whidcy,  1940:  fig.  33).  Both  the  de- 
scription and  the  illustration  were  based 
exclusively  on  the  holotype.  Whitley  re- 
alized that  the  paratypes  were  different 
from  the  holotype  but  considered  them 
to  be  poorly  preserved  specimens  ( Whitley, 
1935:  320). 

A  re-examination  of  the  type  series  re- 
veals that  the  paratypes  are  not  conspecific 
with  the  holotype.  They  are,  instead,  mem- 
bers of  E.  denticulattis.  As  is  shown  in 
Table  25,  counts  and  measurements  from 
the  paratypes  always  fall  within  ranges 
characteristics  of  E.  denticulatus.  Pectoral 
fin  counts,  vertebral  number,  pyloric 
caecum  counts,  and  fin  spine  lengths  are 
particularly  noteworthy  in  this  respect. 
The  paratypes  further  resemble  E.  denti- 
culatus by  bearing  dentigerous  palatines 
and  weak  opercular  armor,   E.  lenimen,  on 


the  other  hand,  is  characterized  by  eden- 
tulous palatines  and  pungent  opercular 
spines. 

Confusion  over  the  type  locality  stems 
from  Whitley's  1940  paper.  The  locality 
is  cited  as  "from  190-320  fathoms,  S.W. 
from  Eucla,  Great  Australian  Bight  [p. 
420] ."  This  contradicts  the  data  presented 
in  the  original  description:  "Great  Aus- 
ti-alian  Bight;  south  from  Eucla,  350  to  450 
fathoms  [p.  231]."  The  1940  citation  is 
extremely  similar  to  station  data  listed  for 
paratypes  AM  E3581-3582  in  1935  ("Great 
Australian  Bight;  SW  from  Eucla,  190-320 
fathoms.  126°  451/2'E  long.  [p.  231]").  In 
the  absence  of  other  information,  it  must  be 
concluded  that  erroneous  locality  data  were 
inserted  in  the  1940  publication  through  an 
editorial  oversight. 

The  most  recent  taxonomic  questions 
arise  from  Whitley's  check  list  of  New 
Zealand  fishes  ( 1968 ) .  This  work  includes 
two  incorrect  citations  in  the  synonymy 
of  E.  lenimen.  The  first  is  based  on  a  fish 
taken  off  the  Chatham  Islands  and  tenta- 
tively identified  as  Hynnodiis  atherinoides 
(Moreland,  1957).  This  specimen  was  later 
re-identified  as  Grahamichthijs  radiatus 
(Moreland,  personal  communication).  The 
second  misidentified  .specimen  is  a  "Big- 
eyed  Cardinal  Fish"  captured  off  Cape 
Palliser,  New  Zealand  (Anonymous,  1961). 
This  fish  is  actually  a  specimen  of  E. 
telescopus  and  is  presently  in  the  collec- 


Ei'ic.oNus  Systematics  •  Mayer       197 


tions  of  the  Dominion  Museum  (DM  3072, 
examined ) . 

Common  names.  None. 

Epigonus  crassicaudus  de  Buen,  1959 
Figure  23 

Epiguinis  crcis.'iicattdii.s  de  Buen,  1959:  196 
(original  description;  preabysnial  zone  off 
Valparaiso,  Chile;  holotype  not  examined,  EBM 
10.183). 

Diagnosis.  E.  crassicaudus  is  strongly 
compressed.  It  reaches  260-270  mm  SL 
and  is  the  second  largest  species  in  the 
genus.  E.  crassicaudus  may  be  distin- 
guished from  E.  trewavasae,  E.  pectinifer, 
E.  rohustus,  and  E.  lenimen  by  its  deep 
head  (18.9-21.2%  SL)  and  deep  body 
(24.3-32.0%  SL).  It  differs  from  remain- 
ing congeners  by  bearing  9  rays  in  the 
second  dorsal  fin  and  6-7  pyloric  caeca. 

Description.  Meristic  values  presented 
in  Table  26;  regression  data  for  morpho- 
metric  traits  presented  in  Table  27. 

Body  elongate,  compressed;  anterodorsal 
profile  rising  from  tip  of  snout  to  occiput, 
becoming  moderately  convex  from  occiput 
to  base  of  first  dorsal  fin.  Body  deep,  24.3- 
32.07f  SL;  caudal  peduncle  broad,  moderate 
to  short,  21.6-26.4%  SL. 

Head  long,  deep,  postorbital  portion 
greatly  expanded,  length  36.8-41.9%  SL; 
height  18.9-21.27r  SL;  snout  moderately 
pointed  in  small  specimens,  blunt  in  adults; 
angle  of  gape  moderate  to  small;  mandible 
long,  strongly  protuberant,  young  bearing 
two  weak  nubs  on  anterior  surface  of  lower 
jaw.  Maxilla  reaching  %-%  eye  length; 
posterior  margin  of  maxilla  broad,  rounded 
or  bearing  posteriormost  point  between 
midline  and  ventral  surface  of  bone.  Eye 
round,  small,  34.2-39.6%  IlL;  surrounded 
by  numerous  small  scale  pockets;  antero- 
dorsal rim  of  orbit  reaching  dorsal  profile^; 
interorbital  region  narrow,  6.2-8.5%    SL. 

Teeth  small,  conical,  occasionally  villi- 
form,  larger  in  small  specimens;  premax- 
illary  teeth  arranged  in  irregular  single 
or    double    rows    tapering    to    single    row 


posteriorly  and  covering  from  %  to  entire 
length  of  bone;  mandibular  teeth  arranged 
in  multiple  rows,  tapering  to  single  row 
posteriorly,  covering  from  V2  to  entire 
IcMigtli  of  dentary;  vomer  edentulous  or 
bearing  up  to  six  irregular  rows  of  minute 
teeth;  palatines  edentulous  or  bearing  1-3 
teeth  anteriorly;  tongue  edentulous. 

Opercular  spine  pungent,  bony,  ventral 
to  3-5  flat,  horny  spinelets;  spine  separated 
from  spinelets  by  narrow  gap;  spinelets 
often  obscured  by  underlying  membranes. 
Preopercular  angle  slightly  produced,  pos- 
terior and/ or  venti'al  surfaces  serrated; 
subopercles  and  interopercles  serrated. 
Gill  rakers  awl-like,  short;  gill  filaments 
long. 

First  dorsal  fin  VII  (22);  second  dorsal 
fin  1,9  (20),  1,10  (2);  anal  fin  II.8  (1), 
11,9  (21);  DJ  2.0-3.6%  SL;  DJ  9.-8-13.2% 
SL;  All  10.3-14.0%  SL;  PJ  13.0-15.5% 
SL. 

Vertebrae  11  +  14  (25);  epipleural  ribs 
6  (2),  7  (16),  inserting  on  vertebrae  1-6 
or  1-7  respectively;  pleural  ribs  9  (25), 
inserting  on  vertebrae  3-11. 

Color  in  alcohol  variable  with  preserva- 
tion; skin  frequently  abraded,  exposing 
underlying  pink  tissue  and  orange-rust  fat 
deposits;  skin  exti'emely  oily;  fin  membranes 
black;  scale  pockets  mottled  with  numerous 
black  melanophores;  dorsal  portion  of  body 
darker  than  ventral;  forehead,  snout,  an- 
terior half  of  mandible,  and  circumorbital 
area  heavily  invested  with  black  pigment; 
opercles  black  or  slate  gray.  Guanine  de- 
posits occasionally  on  opercles,  isthmus, 
pectoral  and  pelvic  fin  bases,  and  al)do- 
men  to  anal  fin;  iris  variable — black,  siher, 
or  black  with  silver  highlights;  mouth  and 
branchial  region  light,  darkening  with  age. 

Description  based  on  27  specimens  80.3- 
262.5  mm  SL. 

Ontogenetic  change.  Two  juvenile  E. 
crassicaudus  (12.2  mm  SL,  MCZ  48857, 
and  21.7  mm  SL,  MCZ  48858)  were  taken 
off  the  Chilean  coast  by  midwater  trawl. 
Although  these  forms  bear  characteristics 
diagnostic  of  the  species,  they  differ  con- 


198       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Table  26.     Epigonus  crassicaudus  meristic  data.    X  =  mean;  SD 

STANfDARD  DEVIATION;   n   =   NUMBER  OF   SPECIMENS. 


X 


Range 


SD 


Pectoral    fin    rays  18.05  17-19  0.58  22 

Gill    rakers  32.27  31-34  0.70  22 

Lateral   line    scales  47.86  46-49  0.85  21 

Pyloric    caeca  6.87  6-     7  0.35  15 


Table  27.     Epigonus  crassicaudus  regression  data,    b   =  regression 

COEFFICIENT  ±  95%  CONFIDENCE  INTERVAL;   a  =   Y  INTERCEPT;   n  =   NUMBER 
OF  SPECIMENS.    AlL  REGRESSIONS  ON  SL. 


b 

a 

n 

HL 

0. 

39 

+ 

0. 

02 

0. 

47 

20 

Body    depth 

0. 

30 

+ 

0. 

04 

-3. 

75 

21 

Head    height 

0. 

21 

+ 

0. 

02 

-1. 

62 

18 

Eye    diameter 

0. 

14 

+ 

0. 

01 

0. 

74 

21 

Snout    length 

0. 

08 

+ 

0. 

01 

0. 

08 

19 

Interorbital    width 

NONLINEAR 

Maxillary    length 

0, 

17 

+ 

0. 

01 

-0. 

49 

20 

Lower    jaw    length 

0. 

20 

+ 

0, 

01 

-1, 

17 

22 

Caudal   peduncle    dep 

th 

0, 

12 

+ 

0. 

01 

-  0, 

65 

20 

Caudal   peduncle    len 

gth 

0. 

22 

+ 

0, 

03 

4, 

32 

20 

D2  I 

0. 

10 

+ 

0. 

02 

3. 

34 

12 

All 

0. 

11 

+ 

0. 

02 

2. 

63 

17 

P2I 

0. 

14 

+ 

0. 

01 

0. 

05 

16 

sideral)ly   in   appearance   and   habit   from 
adnlts. 

Most  .striking  is  the  juvenile  pigment 
pattern.  Pelagic  specimens  are  basically 
pale  yellow  with  large,  brown  patches 
covering  most  of  the  caudal  peduncle. 
Caudal  peduncle  rings,  like  those  found  on 
E.  pandionis  young,  are  absent,  although 
myotomes    are    outlined    by    thin    brown 


bands.  Brown  pigment  extends  anteriorly 
as  a  band  from  the  caudal  peduncle  to  the 
frontal  region  of  the  head.  A  poorly  de- 
fined black  stripe  extends  across  the  snout 
to  the  anterior  rim  of  the  orbit.  In  general, 
juvenile  E.  crassicaudus  resemble  E.  teles- 
copiis  young  figured  by  Koefoed  (1952: 
plate  IIA). 

The  midwater  capture  of  E.  crassicaudus 


Epigonus  Systematics  •  Mayer       199 


juveniles  suggests  that  tlie  life  cycle  of  tlK> 
specit^s  includes  a  pelagic  stage.  Unfortu- 
nately, the  data  available  are  not  sufficient 
to  determine  the  duration  of  this  stage. 

Distribution.  E.  crassicaudus  is  endemic 
to  the  waters  off  central  Chile  (Fig.  21). 
Adults  have  been  captured  by  bottom 
trawls  made  between  200  and  400  meters; 
juveniles  were  taken  by  midwater  trawls 
fishing  from  200  to  270  meters. 

Common  mimes.    None. 

Species  Incertae  Sedis 

Micwichtlnjs  coccoi  Riippell,  1852:  1  (original 
description;  "Mare  siculum";  holotype  not  ex- 
amined, SMF  1069). 

The  original  description  of  M.  coccoi 
provides  only  a  superficial  account  of  the 
holotype.  Subsequent  papers  either  para- 
phrase Riippell's  work  {e.g.,  Canestrini, 
1860;  Doderlein,  1889)  or  are  based  on 
material  not  compared  to  the  holotype 
(i.e.,  Facciola,  1900;  Caporaicco,  1926; 
Gonzales,  1946).  It  is  questionable  whether 
the  latter  specimens  are  conspecific  with 
the  holotype. 

Most  recent  revisers  (e.g.,  Schultz,  1940; 
Norman,  1957)  have  synonymized  Micro- 
ichthys  with  Apogon;  however,  the  data 
are  inconclusive  and  also  suggest  an  affinity 
with  Epigonus  (Eraser,  1972:  5).  A  re- 
examination of  the  holotype  must  be  under- 
taken to   clarify  the  status   of   M.   coccoi. 

A  second  species  of  Microichthys — M. 
sonzoi  Sparta,  1950 — does  not  appear  to  be 
an  Epigonus  on  the  basis  of  vertebral  and 
dorsal  fin  counts.  The  only  known  speci- 
men of  this  species  has  been  lost  (Torton- 
ese,  personal  communication). 

ACKNOWLEDGMENTS 

This  work  would  not  have  been  possible 
without  the  assistance  and  support  of 
niunerous  people.  I  wish  to  thank  the 
following  scientists  and  institutions  for 
material  used  in  this  study:  J.  R.  Paxton 
and  D.  Hoese,  Australian  Museum;  A.  W. 
Wheeler  and  G.   Palmer,   British  Museum 


(Natural  History);  J.  Randall,  Bemice  P. 
Bishop  Museum;  W.  Eschmeyer,  California 
Academy  of  Sciences;  E.  Bertclsen,  Carls- 
bergfondets;  J.  Moreland,  Dominion  Mu- 
seum; L.  P.  Woods,  Eield  Museum  of 
Natural  History;  P.  Struhsaker,  National 
Marine  Fisheries  Service,  Honolulu;  R. 
Raymond,  Instituto  de  Fomento  Pesquero; 
X.  Missonne,  Institut  Royal  des  Sciences 
Naturelles  de  Belgique;  G.  Krefft,  Institut 
fiir  Seefischerei;  M.  M.  Smith,  J.  L.  B. 
Smith  Institute  of  Ichthyology;  I.  Naka- 
mura,  Kyoto  University;  R.  J.  Lavenberg, 
Los  Angeles  County  Museum  of  Natural 
History;  M.  Bauchot,  Museum  National 
d'Histoire  Naturelle;  M.  Poll,  Musee  Royal 
de  I'Afrique  Centrale;  E.  A.  Lachner  and 
T.  H.  Eraser,  National  Museum  of  Natural 
History;  M.-L.  Penrith,  South  African  Mu- 
seum; George  R.  Vliller,  Tropical  Atlantic 
Biological  Laboratory;  R.  S.  Gaille,  Texas 
Parks  and  Wildlife  Department;  M.  Leible, 
Universidad  Catolica  de  Chile;  C.  R. 
Robins,  University  of  Miami;  J.  Nielsen, 
Universitetets  Zoologiske  Museum;  B. 
Nafpaktitis,  University  of  Southern  Cali- 
fornia; T.  Abe,  University  of  Tokyo;  R. 
Backus  and  J.  Craddock,  Woods  Hole 
Oceanographic  Institution;  and  C.  Karrer, 
Zoologisches  Museum,  Berlin.  W.  Klause- 
witz  of  the  Natur-Museimi  Senckenberg 
provided  invaluable  information  on  the 
holotype  of  Microichthys  coccoi,  and  E. 
Tortonese  of  the  Museo  Civico  di  Storia 
Naturale,  Genoa,  answered  numerous  ques- 
tions about  problematical  forms  such  as 
Pomatomichthys  constanciae  and  Micro- 
icJithys  sanzoi. 

I  am  greatly  indebted  to  Ernst  Mayr, 
Giles  W.  Mead  and  Karel  F.  Liem  for 
their  guidance,  criticism,  and  support  of 
my  work.  I  am  also  grateful  to  Richard  L. 
Haedrich  for  reading  the  manuscript  and 
assisting  in  the  planning  of  this  research. 
Special  thanks  are  extended  to  G.  S.  Myers 
for  assistance  with  taxonomic  problems. 

I  wish  to  thank  the  staffs  of  the  Fish 
Department,  Museum  of  Comparative 
Zoology,  and  Department  of  Natinal   Sci- 


200       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


ences,  Boston  University,  for  their  useful 
comments  and  practical  help.  Karen  Green- 
leaf  and  Pat  Allen  typed  the  final  draft  of 
this  manuscript. 

Illustrations  of  eleven  of  the  twelve 
species  of  Epigonus  were  prepared  by  L. 
Laszlo  Meszoly.  Jordan  and  Jordan's  il- 
lustration of  E.  fragilis  (Fig.  10)  was  made 
available  through  the  courtesy  of  the 
Carnegie  Museum. 

Finally,  a  hearty  vielen  Dank  to  my  wife 
for  her  patience,  encouragement,  and  ed- 
itorial assistance. 

Support  for  this  work  was  provided  by 
NSF  Graduate  Fellowships  during  1966 
to  1971  and  a  grant  from  Harvard  Uni- 
versity's Committee  on  Evolutionary  Bi- 
ology (GB7346). 

LITERATURE  CITED 

Anonymous.     1961.     Caught  only  3  times  before. 

Wellington  Evening  Post. 
Bailey,     N.     T.     1959.     Statistical     Methods     in 

Biology.    New   York:    John   Wiley    and    Sons, 

Inc.  200  pp. 
Barnard,    K.    H.     1927.     A    monograph    of    the 

marine  fishes  of  South  Africa,   Part   II.   Ann. 

South    African    Mus.,    12(2):    419-1065. 
Bauchot,  M.  L.,  and  M.  Blanc.     1961.     Poissons 

marins     de     I'Est     Atlantique     Tropical,     II. 

Percoidei      (Teleosteens      Perciformes)      lere 

partie.    Atlantide  Rep.  No.  6:  65-100. 
Bertolini,  F.      1933.     Apogonidae.  Fauna  e  flora 

del    Golfo    di    Napoli.     Uova,    larve    e    stadi 

giovanili    di    Teleostei.      Pubbl.     Staz.     Zool. 

Napoli,  38:  306-309. 
BiNi,    G.      1968.     Atlante    dei    Pesci    delle    Coste 

Italiane,    Vol.    IV.    Roma:    Mondo    Sommerso 

Editrice.  163  pp. 
Bleeker,   p.     1876.     Systema   Percarum   revisum. 

Pars  I  et  II.    Arch.  Neerl.  Sci.,  11:  274-340. 
Braxjer,  a.      1906.     Die  Tiefsee-Fische.    I.  Syste- 

matischer   Teil.    Wiss.    Ergeb.    Deut.    Tiefsee- 

Exped.  "Valdivia,"  1898-1899,  15:   1-420. 
Breder,  C.  M.,  and  D.  E.  Rosen.     1966.     Modes 

of    Reproduction    in    Fishes.      Garden    City, 

N.  Y.:  Natural  History  Press.  941  pp. 
Brown,    M.    E.      1957.     Experimental    studies    on 

growtli.     In   The   Physiology   of    Fishes,    Vol. 

I,  M.  E.  Brown  (ed. ).    New  York:  Academic 

Press,  Inc.,  pp.  361-400. 
Buen,    F.    de.      1959.     Notas    preliminares    sobre 

la    fauna    marina    preabismal    de    Chile,    con 

descripcion    de    una    familia    de    rayas,    dos 


generos   y   siete   especies    nuevos.     Bol.    Mus. 

Nac.    Hist.    Natur.,    27(3):     171-201. 
Canestrini,    J.      1860.     Zur    Systematik    der    Per- 

coiden.     Verb.    Zool.    Bot.    Ver.    Wien,    10: 

291-314. 
Capello,     F.     de     B.     1868.     Peixes     novos     de 

Portugal  e   da  Africa   occidental  e   caracteres 

distinctivos    d'outras    especies    ja    conhecidas. 

J.    Sci.    Math.    Phys.    Natur.   Acad.    Real    Sci. 

Lisboa,   1(2):    154-169. 
Caporaicco,  L.  dl     1926.     II  cranio  de  "Micro- 

ichthvs     coccoi"     Riipp.     Monit.     Zool.     Ital., 

37(6):    127-132. 
Cligny,  a.     1903.     Poissons  des  Cotes  d'Espagne 

et  de  Portugal  (  Ocean  Atlantique ) .    Premiere 

partie.   Boulogne.   30  pp. 
Cocco,   A.     1829.     Su   di  alcuni  nuovi  pesci   de' 

mari  di  Messina.  Giom.  Sci.  Lett.  Art.   Sicil., 

7(77):    138-147. 
.     1885.     Indice    Ittiologico    del    Mare    di 

Messina.    Nat.    Sicil.,    4(4):    85-88. 
CuvrER,   G.     1828.     Des  pomatomes.   In   Histoire 

Naturelle    des    Poissons,    Tome    Second,    G. 

Cuvier    and    A.    Valenciennes.    Paris:    F.    G. 

Levrault,  pp.  169-174. 
Dieuzeide,  R.     1950.     Sur  un  Epigonus  nouveau 

de    la    Mediterranee    (Epigonus    denticulatus, 

nov.  sp.).    Bull.  Sta.  Aquic.  Peche  CastigUone, 

N.S.,  No.  2:  89-105. 
,    M.    Novella,    and    J.    Roland.     1953. 

Catalogue  des  poissons  des  Cotes  algeriennes. 

II.   Osteopterygiens.     Bull.   Sta.   Aquic.   Peche 

Castiglione,  N.S.,  No.  5:  1-258. 
Dlxon,    W.    J.    (ed.).     1967.     BMD    Biomedical 

Computer  Programs.  Univ.  Calif.  Publ.  Auto- 
matic  Computation    No.   2.     Berkeley:    Univ. 

Cahf.  Press.  600  pp. 
DoDERLEiN,    p.     1889.     Manuale    Ittiologico    del 

Mediterraneo.  Fascicolo  IV.  Palermo:  Giornali 

di  Sicilia.  188  pp. 
Dons,   C.     1938.     Notes  on  fishes   III.   Epigonus 

telescopiis     (Risso),    new    to    Norway.      Det 

Kgl.    Nor.    Vidensk.    Selsk.    Forh.,     11(35): 

141-142. 
Eblna,   K.     1931.     Buccal   incubation   in  the   two 

sexes  of  a  percoid  fish,  Apogon  semiUneatus 

T.  &  S.     T.   Imp.  Fish.   Inst.  Tokyo,  27(1): 

19-21. 
Ehrenbaum,  E.     1928.     Rare  fishes  in  the  North 

Sea.    Nature    (London),    121(3053):    709. 
Ekman,     S.     1953.     Zoogeography    of    the     Sea. 

London:    Sidgwick  and  Jackson,  Ltd.  417  pp. 
Facciola,     L.      1900.     Sul     Microichthys     coccoi 

Rapp.  Monit.  Zool.  Ital.,   11(5):    188-194. 
Fowler,   H.   W.     1928.     The  fishes   of  Oceania. 

Mem.  Bernice  P.  Bishop  Mus.,  10:   1-540. 
.      1935.     South     African     fishes     received 

from   Mr.   H.  W.   Bell-Marley  in   1935.    Proc. 

Acad.  Nat.  Sci.  Philadelphia,  87:  361-408. 
.     1936.     The  marine  fishes  of  West  Africa 


Ei'iGONus  Systematics  •  Mayer       201 


based    on    tlie    collection    of    the    American 
Musemn  Congo  Expedition,  1909-1915.    Bull. 
Amer.  Mus.  Nat.  Hist.,   70(2):   607-1493. 
— ,   AND   B.   A.   Bean.      1930.     The   fishes   of 


the  British  Museum,  Vol.  I.  London.  .524  pp. 
1868.     Report  on  a  c(jllection  of  fishes 


the  families  Amiidae,  Chandidae,  Duleidae, 
and  Serranidae  obtained  by  the  United  States 
Bureau  of  Fisheries  Steamer  "Albatross"  in 
1907  to  1910,  chiefly  in  the  Philippine 
Islands  and  adjacent  seas.  Bull.  U.S.  Nat. 
Nhis.,    10(100):    1-334. 

FuASER,  T.  H.  1972.  Comparative  osteology  of 
the  shallow  water  cardinal  fishes  IPerci- 
formes:  Apogonidae]  with  reference  to  the 
s>'steniatics  and  exolution  of  the  family. 
Ichthvol.  Bull.  J.  L.  B.  Smith  Inst.  Ichthyol., 
Rhodes  Univ.,  No.  34:  1-105. 

FucJLLSTER,  F.  C.  1960.  Atlantic  Ocean  atlas  of 
temperature  and  salinity  profiles  and  data 
from  the  International  Geophysical  Year  of 
1957-1958.  Woods  Hole  Oceanogr.  Inst. 
Atlas  Ser.,  1 :  1-209. 

Gall,  J-  le.  1931.  Epigomts  telescopus  Risso 
1810.  In  Faune  Ichthyologique.  Copenhagen, 
Conseil  Permanent  International  pour  1'  Ex- 
ploration   de    la    Mer,    2    pp. 

GiGLiOLi,  E.  H.  1880.  Elenco  dei  Mammiferi, 
degli  Uccelli  e  dei  Rettili  Ittiofagi  Ap- 
partenenti  alia  Fauna  Italica  e  Catalogo  degli 
Anfibi  e  dei  Pesci  Italiani.  Firenze:  Staniperia 
Reale.  85  pp. 

Gilbert,  C.  H.  1905.  The  deep-sea  fishes  of 
the  Hawaiian  Islands.  In  The  Aquatic  Re- 
sources of  the  Hawaiian  Islands.  II,  D.  S. 
Jordan  and  B.  W.  Evermann  (eds. ).  Bull. 
U.  S.  Fish.  Comm.,  23:  575-716. 

Gilchrist,  J.  D.  F.  1922.  Report  No.  2  for  the 
year  1921.  Fish.  Mar.  Biol.  Surv.  South 
Africa,  Rep.  No.  2:  1-84. 

,    AND    C.    vox    BoxDE.     1924.     Deep-sea 

fishes  procured  by  the  S.  S.  "Pickle"  (Part 
II).  Fish.  Mar.  Biol.  Surv.  South  Africa, 
Rep.  No.  3,  Spec.  Rep.  VII:    1-24. 

GoxzALES,  T.  1946.  Contributo  alia  conoscenza 
dello  s\iluppo  post-embrionale  in  Micro- 
ichtJit/s  coccoi,  Riippell.  Boll.  Pesca  Piscicolt. 
Idrobiol.,  Ser.  22,  1  ( 1 ) :  39-46. 

GooDE,  G.  B.,  AXD  T.  H.  Beax.  1881.  De- 
scription of  a  new  species  of  fish,  Apogon 
))andionis,  from  deep  water  off  the  mouth 
of  Chesapeake  Bav.  Proc.  U.  S.  Nat.  Mus., 
4:    160-161. 

,  AXD  .  1896.  Oceanic  Ichthy- 
ology. Wa.shingtou,  D.  C:  Smithsonian  In- 
stitution. 553  pp. 

GosLixE,  \V.  A.  1961.  The  perciform  caudal 
skeleton.     Copeia,    1961(3):    265-270. 

,   AXD  \'.   E.   Brock.      1960.     Handbook   of 

Hawaiian  Fishes.  Honolulu:  Univ.  Hawaii 
Press.  372  pp. 

Giinther,   A.      1859.     Catalogue   of   the   Fishes    in 


made  at  St.  Helena  by  J.  C.  Melli.ss,  Esq. 
Proc.  Zool.  Soc.  London,  pp.  22.5-228. 

Haxeda,  Y.,  F.  H.  Johxsox,  and  O.  Shi.mo.mura. 
1966.  The  origin  of  luciferin  in  the  luminous 
ducts  of  Pampriacmithus  ransonncti,  Pem- 
plicris  kltiiizingcii,  and  Apogon  cllioti.  In 
Bioluminescence  in  Progress,  F.  H.  Johnson 
and  Y.  Haneda  (eds.).  Princeton:  Princeton 
Univ.  Press,  pp.  533-545. 

Holt,  E.  W.,  AND  W.  L.  Calderwood.  1895. 
Survey  of  fishing  grounds,  west  coast  of 
Ireland,  1890-1891.  Report  on  the  rarer 
fishes.  Scient.  Trans.  R.  Dublin  Soc,  Ser. 
2,   5:    361-524. 

HuHBS,  C.  L.,  AND  K.  F.  Lacler.  1958.  Fishes 
of  the  Great  Lakes  Region.  Ann  Arbor: 
Univ.  Michigan  Press.  213  pp. 

I\\Ai,  T.  1959.  Notes  on  the  luminous  organ 
of  the  apogonid  fish,  Siphaiiiia  incijiinoi. 
Ann.  Mag.  Natur.  Hist.,  Ser.  13,  2(21): 
545-550. 

Jordan,  D.  S.  1917.  Notes  on  Glossamia  and 
related  genera  of  cardinal  fishes.  Copeia, 
No.  44:  46-47. 

,    AND    C.    H.    Gilbert.     1882.     Synopsis 

of  the  fishes  of  North  America.  Bull.  U.  S. 
Nat.   Mus.,   16:    1-1018. 

,    AND    E.    K.    Jordan.     1922.     A    hst    of 

the  fishes  of  Hawaii,  with  notes  and  de- 
scriptions of  new  species.  Mem.  Carnegie 
Mus.,   10(1):    1-92. 

Kamohara,  T.  1952.  Revised  descriptions  of 
the  offshore  bottom-fishes  of  Prov.  Tosa, 
Shikoku,  Japan.  Rep.  Kochi  Univ.  Nat.  Sci., 
No.  3:  1-22. 

Koefoed,  E.  1952.  Zeomorphi,  Percomorphi, 
Plectognathi.  Rep.  Sci.  Res.  Michael  Sars 
N.  Atl.  Deep-Sea  Exped.  1910,  4,  Pt.  2(2): 
1-26. 

Lowe,  R.  T.  1841.  A  synopsis  of  the  fishes  of 
Madeira;  with  principal  synonyms,  Portu- 
guese names,  and  characters  of  the  new 
genera  and  species.  Trans.  Zool.  Soc.  London, 
2(14):    173-200. 

LozANO,  L.  1934.  Algunos  peces  pelagicos  o 
de  profundidad  procedentes  del  Mediterraneo 
occidental.  Bol.  Soc.  E.span.  Hist.  Natur., 
34:   85-92. 

Matsubara,  K.  1936.  Biometry  of  two  species 
of  Japanese  cardinal-fishes,  with  special 
reference  to  their  taxononn.  J.  Imp.  Fish. 
In.st.  Tokyo,  31:    119-1.30. 

Maurin,  C.  1968.  Ecologie  ichthyologiques  des 
fonds  Chalutables  atlantiques  (de  la  baie 
ibero-marocaine  a  la  Mamitanie)  et  de  la 
Mediterranee  occidentale.  Theses  presentees 
a  la  Faculte  des  Sciences  de  I'Universite  de 
Nancy.    No.  d'Ordre:  A.  O.  2  182.  145  pp. 


202       Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  3 


Mayer,  G.  F.  1972.  Systematics,  functional 
anatomy,  and  ecology  of  the  caidinalfish 
genus  Epigonus  ( Apogonidae ) .  Ph.  D. 
Thesis,  Hanard  University.    190  pp. 

Mayr,  E.  1969.  Principles  of  Systematic  Zo- 
ology.   New  York:  McGraw-Hill,  Inc.  428  pp. 

Mead,  G.  W.,  and  M.  G.  Bradbury.  1963. 
Names  of  bones.  In  Fishes  of  the  Western 
North  Atlantic,  H.  B.  Bigelow  and  W.  C. 
Schroeder  (eds.).  Mem.  Sears  Found.  Mar. 
Res.,  1(3):  20-23. 

MoNOD,  T.  1968.  Le  complexe  urophore  des 
poissons  teleosteens.  Mem.  Inst.  Fondam. 
Afr.  Noire,  No.  81.  705  pp. 

MoREAU,  E.  1881.  Histoire  Naturelle  des  Pois- 
sons de  la  France.  Vol.  II.  Paris:  G.  Masson. 
572  pp. 

MoRELAND,  J.  1957.  Appendix  6:  Report  on 
the  fishes.  New  Zealand  Dep.  Sci.  Ind.  Res. 
Bull.,   122:  34. 

Navarro,  F.  de  P.  1942.  Nota  preliminar  sobre 
los  paces  de  la  costa  de  Africa,  desde  el  Cabo 
Bojador  a  la  Bahia  de  Tanit  ( Resultados  de 
una  compana  industrial  de  pesca  d'arrastre). 
Bol.  Real  Soc.  Espan.  Hist.  Natur.,  40(5-6): 
189-214. 

,    F.    LozANO,    J.    M.    NovAz,    E.    Otero, 

AND  J.  Sainz  Pardo.  1943.  La  pesca  d' 
arrastre  en  los  fondos  del  Cabo  Blanco  y 
del  Banco  Arguin  (Africa  Sahariana).  Trab. 
Inst.  Espaii.  Oceanogr.  Numero  18.  225  pp. 

Nohre,  a.  1935.  Descricao  dos  Peixes  de 
Portugal.  Vertebrados  (mamiferos,  reptis  e 
peixes).  I.  Fauna  Marinha  de  Portugal. 
Porto.  574  pp. 

Norman,  J.  R.  1939.  Fishes.  Sci.  Rep.  John 
Murray  Exped.,  7(1):    1-116. 

.      1957.     A  Draft  Synopsis  of  the  Orders, 

Families  and  Genera  of  Recent  Fish  and 
Fish-like  Vertebrates.  London:  British  Mu- 
seum (Natural  History).  649  pp. 

Nowlin,  W.  D.,  jr.,  and  H.  J.  McLellan. 
1967.  A  characterization  of  the  Gulf  of 
Mexico  waters  in  winter.  ].  Mar.  Res.,  25(1): 
29-59. 

Osomo,  B.  1898.  Da  distribviicao  geographica 
dos  peixes  e  crustaceos  colhidos  nas  pos- 
sessoes  Portuguezos  d'Afrique  occidental  e 
existentes  no  Museu  Nacional  de  Lisboa. 
J.  Sci.  Math.  Phys.  Natur.  Acad.  Real  Sci. 
Lisboa,  Ser.  2,   5:    185-202. 

PiETSCHMANN,  V.  1930.  Rcuiarks  on  Pacific 
fishes.  Bull.  Bernice  P.  Bi.shop  Mus.,  73: 
1-24. 

Poll,  M.  1954.  Poissons  IV.  Teleosteens 
acanthopterygiens  (premiere  partie).  Re- 
sidtats  scientifiques.  Expedition  Oceano- 
graphique  Beige  dans  les  Eaux  Cotieres 
Africaines  de  I'Atlantique  Sud  (1918-1949), 
4(3A):   1-390. 


Radcliffe,  L.  1912.  Descriptions  of  fifteen 
new  fishes  of  the  family  Gheilodipteridae, 
from  the  Pliilippine  Islands  and  contiguous 
waters.  Proc.  U.  S.  Nat.  Mus.,  41(1868): 
431-446. 

Rafinesque,  G.  S.  1810.  Indice  d'  Ittiologia 
Siciliana;  Ossia,  Gatalogo  Metodico  dei  Nomi 
Latini  Italiani,  e  Siciliani  dei  Pesci,  che  si 
Rinvengono  in  Sicilia:  Disposti  Secondo  un 
Metodo  Naturale  e  Seguito  da  un  Appendice 
che  Gontiene  la  Descrizione  di  Alcuni  Nuovi 
Pesci  Siciliani.  Messina:  Presso  Giovanni 
del  Nobolo.  70  pp. 

Risso,  A.  1810.  Ichthyologie  de  Nice  ou  Histoire 
naturelle  des  poissons  du  department  des 
Alpes  Maritimes.  Paris:  Chez  F.  Schoell. 
388  pp. 

Royce,  W.  R.  1957.  Statistical  comparison  of 
morphological  data.  In  Contributions  to  the 
Study  of  Subpopulations  of  Fishes,  J.  G. 
Marr  (coordinator).  U.  S.  Fish.  Wildl.  Serv. 
Spec.  Sci.  Rep.  Fish.,  No.  208:  7-28. 

RuppELL,  W.  1852.  \'erzeichniss  der  in  dem 
Museum  der  Senckenbergisclien  naturfor- 
schenden  Gesellschaft  aufgestellten  Samm- 
lungen.  Vierte  Abteilung.  Fische  und  deren 
Skelette.  Frankfurt-a.M.:  J.  D.  Sauerliinder. 
40  pp. 

Sakamoto,  K.  1930.  Buccal  incubation  in  the 
percoid  fish,  Apogon  lineatus  T.  &  S.  J.  Imp. 
Fish.  Inst.  Tokyo,  26(  1 ) :  9-10. 

ScHULTZ,  L.  p.  1940.  Two  new  genera  and 
three  new  species  of  cheilodipterid  fishes, 
with  notes  on  the  other  genera  of  the  familv. 
Proc.   U.   S.   Nat.   Mus.,  88(3085):   403-423. 

Scott,  T.  D.  1962.  The  Marine  and  Fresh 
Water  Fishes  of  South  Australia.  Adelaide: 
South  Australian  Branch  of  the  British  Sci- 
ence Guild.   338  pp. 

Simpson,  G.  G.,  A.  Roe,  and  R.  C.  Lewontin. 
1960.  Quantitative  Zoology.  New  York:  Har- 
court,  Brace  and  World,  Inc.  440  pp. 

Smith,  G.  L.,  E.  H.  Atz,  and  J.  C.  Tyler.  1971. 
Aspects  of  oral  brooding  in  the  cardinalfish 
Cheilodiptcius  af  finis  Poey  (Apogonidae). 
Amer.   Mus.   Novitates,   No.   2456:    1-11. 

Smith,  J.  L.  B.  1949a.  Forty-two  fishes  new 
to  South  Africa,  with  notes  on  others.  Ann. 
Mag.   Natur.   Hist.,   Ser.    12,   2:    97-111. 

— .      1949b.     The     Sea    Fishes    of    Southern 

Africa.    South  Africa:   Central  News  Agenc>-, 
Ltd.  550  pp. 

1961.     Fishes  of  the  family  Apogonidae 


of  the  western  Indian  Ocean  and  the  Red 
Sea.  Ichth%ol.  Bull.  Rhodes  Univ.,  No.  22: 
373-418. 
Sparta,  A.  1950.  Su  di  una  nuova  specie  di 
Micwichthi/s:  M.  sanzoi  (n.  sp.).  Boll.  Pesca, 
Piscicolt.  Idrobiol.,  Ser.  26,  5(2):  202-206. 


EricoNus  Systematics  •  Mayer       203 


Stein'dachner,  F.  1891.  lehthNologischc  Bei- 
triiKt'  (X\^).  i'her  ciiiige  scltenc  iiiul  ucnc 
Fisc'luiitc'ii  aus  ck'in  caiuiiisclu'ii  Aicliipcl. 
Sitzber.  Akad.  VViss.  Wien,  100(1):  343-374. 

.      1907.     Fi.sche      aiis      Siidaiabicn      imd 

Sokotra.  Denkschr.  Akad.  Wiss.  Wien,  71: 
123-168. 

Taylok,  W.  R.  1967.  An  enz\-matic  method  of 
elearing  and  staining  small  \eitel)iates.  Proe. 
U.S.   Nat.    Mns.,    122(3596):    1-17. 

Tinker,  S.  W.  1944.  Hawaiian  Fishes.  Ilono- 
kdii:   Tongg  PnbHshing  Company.    404  pp. 

ToRTONESE,  E.  1952.  Un  percoide  marino  e 
batifilo  nno\o  per  Tittiofauna  itaHana  (Epi- 
^oinis  clciilicuUitiis  Dienz.).  Boll.  Pesca 
Pi.scicolt.   Idrobiol.   Ser.  28,   7(1):   72-74. 

,  AND  L.  C.  QuEiROLO.     1970.     Contribute 

alio  studio  dell  ittiofauna  del  Mar  Ligure 
orientale.  Ann.  Mus.  Civ.  St.  Natur.  Genova, 
78:   21-46. 

Vaillaxt,  L.  1888.  Poissons.  E.xpeditions  Sci- 
entifiques  du  Travailleur  et  du  Talisman 
pendant  les  Annees  1880,  1881,  1882,  1883. 
Paris:  G.  Masson.  406  pp. 

V^ALENCiENNES,  A.  1830.  Additions  et  correc- 
tions anx  tomes  II,  III,  IV,  et  V.  In  Historic 
Naturelle  des  Poissons,  Tome  Sixieme,  G. 
Cuvier  and  A.  Valenciennes.  Paris:  F.  G. 
Levrault,  pp.  495-559. 

.  1837-1844.  Ichthyologie  des  iles  Can- 
aries, on  Histoire  naturelle  des  poissons 
raportes  par  M.  M.  Webb  et  Berthelot.  In 
Histoire  Naturelle  des  lies  Canaries,  P.  B. 
Webb  and  S.  Berthelot  (eds.).  Vol.  2,  Pt.  2. 
Paris,  pp.  1-109.  [pp.  1-8  published  Dec. 
1842;  plate  1  published  May  1837.] 

Weuer,  M.,  and  L.  F.  de  Beaufort.  1929.  The 
Fishes  of  the  Indo-Australian  Archipelago. 
y.  Anacanthini,  Allotriognathi,  Heterosomata, 
Berycomorphi,  Percomorplii:  Families  Kuhli- 
idae,  Apogonidae,  Plesiopidae,  Pseudo- 
plesiopidae,  Priacanthidae,  Centropomidae. 
Leiden:  E.  J.  Brill,  Ltd.  458  pp. 

Whitley,  G.  P.  1935.  Studies  in  ichthyology. 
No.  9.  Rec.  Aust.  Mus.  Sydney,  19(4): 
215-250. 

-.      1940.      Illustrations    of    some    Australian 

fishes.    Aust.    Zool.,    9(4):     397-428. 

.  1968.  A  check-list  of  the  fishes  re- 
corded from  the  New  Zealand  region.  Aust. 
Zool.,    15(1):    1-102. 

Williams,  F.  1968.  Report  on  the  Gnincan 
Trawling    Survey,    \'ol.     I.     General     Report. 


OAU     Scientific,     Technical     and     Research 
('oninn'ssion,   Ful)licali<)n  99.  828  pp. 
W'iisi,    (;.      1961.      Slratitication    and    Circulati(m 
in  the  Antillean-Caribbcan  Basins.    New  York: 
C]()luml)ia  l'ni\ .  Press.  201  pp. 

APPENDIX 

The  following  chart  lists  all  meristic  and 
niorphometric  data  for  the  holotypcs  of 
E.  oUiiolepis  sp.  nov.  and  E.  pectinifer  sp. 
nov.  Measurements  are  given  in  milli- 
meters. 


E. 

oligolepis 

E.     pectinifer 

USNM    2  077  18 

USNM    207725 

MERISTIC     DATA 

Dorsal    fin 

VII-I,  10 

VII  -I,  9 

Anal    fin 

II,  9 

II,  9 

Pectoral    fin 

18 

15 

Pelvic    fin 

I,  5 

I,  5 

Lateral    line 
sc  ale  s 

34 

47 

Gill    r  ake  r  s 

31 

27 

Pyloric    caeca 

10 

6 

Ve  r tebr  ae 

10  +  15 

10  +  15 

Pleural    ribs 

7 

8 

E  p  i  p  1  e  u  r  a  1    rib 

s 

7 

6 

MORPHOMETRIC 

DATA 

SL 

90.  8 

114.  3 

HL 

33.  2 

40.  5 

Body    depth 

21.2 

28.  1 

Head    height 

17.  1 

18.  8 

Eye    diameter 

14.  5 

16.  1 

Snout    1  e  n  g  t  li 

7.  8 

10.  9 

I  n  t  e  r  0  r  b  i  t  a  1    w  i 

idth 

8.  4 

10.  0 

Maxillary    leng 

th 

15.  6 

18.  2 

Lower    jaw    len 

gth 

16.  3 

18.  5 

Caudal    p  c  <l  u  n  c  : 
d  e  p  t  h 

le 

8.  9 

11.  7 

Caudal    p  e  d  u  n  c  : 
length 

le 

23.  6 

32.  1 

D2l 

10.  9 

13.  7 

All 

11.1 

13.  7 

P2I 

12.  3 

14.  4 

us  ISSN  0027-4100 


BulLetln  OF  THE 

Museum   of 

Comparative 

Zoology 


The  Spider  Family  Anyphaenidae 
in  America  North  of  Mexico 


NORMAN  PLATNICK 


HARVARD  UNIVERSITY 

CAMBRIDGE,  MASSACHUSETTS,    U.S.A. 


VOLUME  146,  NUMBER  4 
19  SEPTEMBER  1974 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED  BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 


Breviora  1952- 

BuLLETiN  186a- 

Memoirs  1864-1938 

JoHNSONiA,  Department  of  Mollusks,  1941- 

OccAsiONAL  Papers  on  Mollusks,  1945- 

SPECIAL  PUBLICATIONS. 

1.  Whittington,  H.  B.,  and  E.  D.  I.  Rolfe  (eds.),   1963.    Phylogeny  and 
Evolution  of  Crustacea.  192  pp. 

2.  Turner,  R.  D.,  1966.  A  Survey  and  Illustrated  Catalogue  of  the  Teredini- 
dae  (Mollusca:  Bivalvia).  265  pp. 

3.  Sprinkle,  J.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echinoderms. 
284  pp. 

4.  Eaton,  R.  J.  E.,  1974.  A  Flora  of  Concord.  211  pp. 

Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.    Fishes  of  the  Gulf  of  Maine. 
Reprint. 

Brues,  C.  T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.    Classification  of 
Insects. 

Creighton,  W.  S.,  1950.    The  Ants  of  North  America.    Reprint. 

Lyman,   C.   P.,   and  A.   R.    Dawe    (eds.),    1960.    Symposium   on   Natural 
Mammalian  Hibernation. 

Peters'  Check-list  of  Birds  of  the  World,  vols.  2-7,  9,  10,  12-15. 

Proceedings  of  the  Nevi'  England  Zoological  Club  1899-1948.    (Complete 
sets  only.) 

Publications  of  the  Boston  Society  of  Natural  History. 

Price  list  and  catalog  of  MCZ  publications  may  be  obtained  from  Publications 
Office,  Museum  of  Comparative  Zoology,  Harvard  University,  Cambridge,  Massa- 
chusetts, 02138,  U.S.A. 

©  The  President  and  Fellows  of  Harvard  College  1974. 


THE  SPIDER  FAMILY  ANYPHAENIDAE  IN  AMERICA 
NORTH  OF  MEXICO' 

NORMAN   PLATNICK- 


Abstract.  E\'idence  from  the  tracheal  system, 
claw  tufts  and  courtship  behavior  is  used  to 
justify  the  family  status  of  Anyphaenidae.  Sug- 
gested relationships  between  Anyphaenidae  and 
Clubionidae,  Amaurobiidae  and  Argyronetidae 
are  disclaimed.  The  faniih'  Amaurobioididae  is 
newly  synonymized  with  Anyphaenidae.  Generic 
problems  within  the  family  are  discussed.  The 
thirty-six  species  occurring  north  of  Mexico  are 
described,  their  diagnostic  characters  pointed  out 
and  illustrated,  their  distributions  mapped,  notes 
on  their  habits  given,  and  keys  to  genera, 
species  groups  and  species  provided.  The  genera 
AuiipliaencUa  and  Ciagus  are  newly  synonymized 
witli  Wulfila.  Thirteen  species  are  described  as 
new:  Amjpliacna  aJachua,  A.  arhida,  A.  uutumna, 
A.  catalina,  A.  cochise,  A.  gertschi,  A.  fiiJjIwides, 
A.  hespar,  A.  lacka,  A.  rita,  Aijslia  aninda, 
Wulfila  hnjantac  and  W.  wiinda.  Nineteen  new 
synonymies  are  recognized. 

INTRODUCTION 

This  study  had  three  objectives:  to  de- 
termine whether  or  not  the  anyphaenids 
should  be  treated  as  a  distinct  family;  to 
examine  the  relationships  between  the  any- 
phaenids and  the  other  groups  of  spiders 
with  which  they  have  been  associated  in 
the  past;  and  to  reclassify  the  species  oc- 


'  This  study  was  presented  to  the  Department 
of  Biology  at  Harvard  University  in  partial  ful- 
fillment of  the  requirements  for  the  degree  of 
Doctor  of  Pliilosophy. 

"Present  address:  Department  of  Entomology, 
The  American  Nhiseum  of  Natural  History,  Cen- 
tral Park  West  at  79th  Street,  New  York,  New 
York  10024. 


curring  north  of  Mexico  on  generic  and 
specific  levels. 

The  anyphaenids  are  a  diverse  group 
with  perhaps  five  hundred  species.  Thirty- 
six  species  are  known  to  occur  in  America 
north  of  Mexico  and  are  included  here. 
About  375  species  have  been  described 
from  the  Neotropic  region,  as  well  as 
around  ten  from  the  Palearctic  and  five 
from  the  Oriental.  The  South  American 
species  show  the  widest  spectrum  of  body 
forms;  they  range  from  2-25  mm  in  length 
and  are  often  intricately  colored  or  have 
peculiarly  elongate  chelicerae  or  legs. 

As  in  most  spiders,  little  is  known  of  the 
ecology  or  behavior  of  anyphaenids.  They 
are  wandering  hunters.  In  the  eastern 
United  States,  where  long-legged  species 
predominate,  they  are  most  often  collected 
by  sweeping  foliage  in  fields  and  meadows, 
and  seem  to  be  primarily  noctiu-nal.  How- 
ever, in  the  western  United  States,  where 
most  species  have  shorter  legs,  they  are  usu- 
ally found  in  forests  by  sifting  through 
litter  and  turning  logs  and  stones.  They 
feed  on  various  groups  of  insects,  and 
though  they  have  been  observed  to  prey 
heavily  on  such  Lepidoptera  as  the  fall 
webworm,  IlijpJiantria  cunea  (Warren  et 
al.,  1967),  they  are  proliably  not  very  se- 
lective. In  captivity  they  will  consume 
Drosophila  eagerly.  Their  principal  ene- 
mies in  nature  are  the  mud-dauber  wasps 
of  the  family  Sphecidae,  as  evidenced  from 
the    hundreds    of   individuals,    particularly 


Bull.  Nhis.  Comp.  Zool,  146(4) :  205-266,  September,  1974        205 


206         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


of  the  diurnally  active  genus  Aijsha,  that 
are  frequently  collected  from  wasp  nests. 
Krombein  ( 1967 )  cites  especially  the  wasp 
genus  Trypargilum  in  this  respect.  Like 
most  nearctic  spiders,  males  and  females 
usually  mature  in  early  spring,  with  males 
living  through  early  summer  and  females 
living  through  the  summer.  In  some  south- 
ern species,  however,  both  sexes  are  found 
matm-e  year-round.  Also,  some  species  in 
the  Amjphaena  celer  group  are  matiu-e 
throughout  the  winter.  Anyphaenids  make 
little  use  of  silk,  other  than  in  building  re- 
treats under  leaves  or  stones  and  of  course 
in  building  egg  sacs,  which  are  usually 
round,  made  of  soft  white  silk,  not  leathery 
or  papery,  and  contain  between  50  and 
150  eggs. 

The  North  American  species  are  2-9  mm 
long;  the  largest  species  belong  to  the 
genus  Aijsha,  the  smallest  to  Wulfih. 
There  are  always  eight  eyes  in  two  rows; 
the  median  eyes  are  usually  closer  to  the 
laterals  than  to  each  other;  unlike  many 
gnaphosids,  the  eyes  are  always  round,  and 
unlike  many  clubionids,  the  anterior  me- 
dian eyes  are  usually  smaller  than  the 
others.  Other  than  the  genitalia,  the  main 
structural  differences  between  males  and 
females  are  the  sternal  and  coxal  modifi- 
cations (pointed  spurs,  rounded  knobs,  or 
clumps  of  short  thick  setae)  found  on 
males  in  some  groups. 

In  many  species  groups  it  would  be  im- 
possible to  distinguish  the  species  without 
using  genitalic  characters.  The  palpus 
(Text-fig.  3)  usually  has  a  large  median 
apophysis  (the  shape  of  which  is  often 
species-specific),  a  small  conductor  and  a 
conspicuous  embolus.  Besides  the  retrolat- 
eral  tibial  apophysis  (almost  always  of 
great  diagnostic  value)  a  ventral  tibial 
apophysis  (some  Aijsha)  or  a  retrolateral 
patellar  apophysis  (some  Teudis)  may  be 
present.  The  female  epigyna  and  internal 
genitalia  are  extremely  diverse  and  diffi- 
cult to  characterize.  The  two  epigynal 
openings  are  located  posteriorly  and  are 
extremely  difficult  to  see  unless  a  portion 


of  the  male  embolus  has  been  left  behind 
after  mating.  Many  species  have  an  addi- 
tional anterior  median  epigynal  opening 
into  which  the  retrolateral  tibial  apophysis 
or  median  apophysis  fits  during  mating. 
The  genitalia  of  anyphaenids,  particularly 
of  the  South  American  species,  are  more 
complex  than  those  of  clubionids  and 
gnaphosids.  Among  the  clubionids,  only 
Chiracanthium  has  genitalia  that  seem  in 
any  way  close  to  those  of  anyphaenids. 

For  the  area  treated  here,  only  three  im- 
portant papers  have  been  published  on 
anyphaenids.  Bryant  (1931)  summarized 
the  very  sparse  data  then  available  on  the 
group  in  the  United  States,  while  Chicker- 
ing  (1937,  1940)  described  many  of  the 
species  occurring  in  Panama  and  the  Canal 
Zone,  a  number  of  which  also  occur  in  the 
United  States. 

ACKNOWLEDGEMENTS 

I  would  like  to  thank  first  Herbert  W. 
Levi  for  his  painstaking  and  patient  help 
with  all  aspects  of  this  project.  Willis 
Gertsch  contributed  much  of  his  knowl- 
edge of  the  group  as  well  as  the  drawings, 
done  by  the  late  Wilson  Ivie,  of  the  genus 
Oxijsorna. 

This  investigation  was  supported  in  part 
by  Public  Health  Service  Research  Grant 
AI-01944  from  the  National  Institutes  of 
Allergy  and  Infectious  Diseases,  H.  W. 
Levi,  principal  investigator;  by  Grant  GB- 
36161  from  the  National  Science  Founda- 
tion, H.  W\  Levi,  principal  investigator; 
and  by  Grant  GB-19922  from  the  National 
Science  Foundation,  R.  C.  Rollins,  princi- 
pal investigator.  The  Department  of  Biol- 
ogy, Harvard  University,  by  means  of 
Summer  Research  Grants  in  Evolutionary 
Biology  in  1971  and  1972  and  a  Richmond 
Fellowship  in  1973,  afforded  me  much  of 
the  time  and  field  work  necessary  to  com- 
plete this  work.  Miss  Suzanne  Barbier  of 
Radcliffe  Gollege  assisted  greatly  with  the 
examination  of  tracheal  systems  and  her 
work  is  deeply  appreciated. 

Finally,    the    following    people    loaned 


SpTDKn  FA\rii,v  ANVPiiAEXinAK  •  Platnirk 


207 


specimens  from  their  private  collections 
or  from  their  cited  institntions:  Paul  II. 
Aniaud,  Jr.  (California  Academy  of  Sci- 
ences), Joseph  A.  Beatty,  Jr.,  James  E. 
Carico,  John  A.  L.  Cooke  (American  Mu- 
seum of  Natural  History),  Charles  D. 
Dondale  (Canadian  National  Collections), 
R.  R.  Forster  (Otago  Museum),  Willis  J. 
Gertsch  (American  Museum  of  Natural 
History),  Al  Jung,  B.  J.  Kaston,  Robin 
Leech,  William  B.  Peck,  Vince  Roth,  Rich- 
ard J.  Sauer  (Michigan  State  University), 
William  A.  Shear,  Bea  Vogel,  H.  K.  Wal- 
lace, Fred  R.  Wanless  (British  Museum, 
Natiu-al  History),  and  Howard  V.  Weems 
(Florida  State  Collection  of  Arthropods). 

THE  FAMILY  STATUS  OF 
ANYPHAENIDAE 

Simon  considered  tlie  anyphaenids  to  be 
a  subfamily  of  the  large  family  Clubionidae 
and  used  as  the  key  character  for  distin- 
guishing the  anyphaenids  the  advanced 
placement  of  the  tracheal  spiracle.  Later 
authors,  notably  Petrunkevitch  and  Bris- 
towe,  thought  this  character  so  significant 
that  they  gave  the  anyphaenids  family  sta- 
tus, though  still  believing  the  group  to  be 
closely  related  to  the  Clubionidae.  The 
comparatively  recent  discovery  that  in 
some  families  closely  related,  congeneric 
species  sometimes  have  very  different  res- 
piratory systems  (see  Levi,  1967)  has  led 
most  arachnologists  to  denigrate  the  im- 
portance of  respiratory  structm-es  as  macro- 
taxonomic  characters.  Thus  most  modern 
arachnological  works  still  treat  the  any- 
phaenids as  a  subfamily  of  Clubionidae. 
A  notable  exception,  however,  is  Lehtinen 
( 1967 ) ,  who  maintains  ( correctly,  I  be- 
lieve) that  the  classical  family  Clubionidae 
is  a  highly  polyphyletic  assemblage  of  un- 
related two-clawed  spiders  that  lack  any 
noticeable  modifications  of  the  body.  Leh- 
tinen splits  the  clubionids  into  several  fam- 
ilies, largely  but  not  strictly  along  the  lines 
of  the  old  subfamily  divisions,  and  accords 
the    anyphaenids    full    status    as    a    family. 


Forster  (1970)  agrees  with  this  assessment 
of  the  anyphaenids. 

To  check  on  the  validity  of  this  classifi- 
cation, a  variety  of  clubionid  genera  were 
examined  and  compared  with  anyphaenids, 
with  the  result  that  the  anyphaenids  arc 
here  considered  a  distinct  family,  for  two 
major  reasons.  One  is  the  classical  reason 
— the  tracheal  system.  Examination  of  the 
tracheae  of  males  and  females  of  the  club- 
ionids CAuhiona  o])esa  Hentz,  Chiracan- 
thium  mildei  L.  Koch,  Trachelus  tratujuil- 
his-  (Hentz),  Castkineira  cin<i,uluta  (C.  L. 
Koch),  Agroeca  pratensis  Emerton,  Phniro- 
timpus  alarius  (Hentz),  and  the  any- 
phaenids Am/pJmena  celer  (Hentz),  Amj- 
phaena  pectorosa  L.  Koch,  Amjphaena 
calif  ornica  ( Banks )  and  Aijsha  <i,racilis 
(Hentz)  disclosed  three  major  differences 
between  anyphaenid  and  clubionid  tra- 
cheae (see  Methods  for  the  technique 
used).  First,  anyphaenid  tracheae  extend 
through  the  pedicel  of  the  spider  into  the 
cephalothorax  and  legs,  while  those  of 
clubionids  are  restricted  to  the  abdomen 
(see  Figs.  47  and  50).  Associated  with  this 
is  the  externally  observable  advanced 
placement  of  the  tracheal  spiracle  in  any- 
phaenids. Second,  the  tiacheae  are  rela- 
tively much  larger  in  anyphaenids.  In  all 
the  clubionids  examined,  even  the  main 
tracheal  tubes  are  very  thin  and  narrow; 
anyphaenid  tracheae  are  three  to  four 
times  as  wide.  Third,  none  of  the  clubionid 
species  examined  showed  any  sexual  di- 
morphism in  the  tracheal  system,  whereas 
male  anyphaenids  have  considerably  larger 
tracheae  than  do  the  females.  The  size 
of  the  tracheae  may  be  correlated  with  the 
high  activity  levels  of  anyphaenids:  my 
collecting  experience  indicates  that  they 
can  run  extremely  rapidly  when  disturbed. 

The  larger  size  of  the  tracheae  in  males 
may  be  associated  with  the  increased  respi- 
ration necessary  for  the  extra  activity  re- 
(juired  to  locate,  court  and  copulate  with 
a  female.  Anyphaenid  courtship  is  (.ex- 
tremely active;  films  of  the  courtship  of 
Anijphacna  accentiiata  show  that  the  abdo- 


208         Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  4 


embolus 


median  apophysis 


conductor 


retroloteral  tibial 
apophysis  (RTA) 


Text-Figures  1-3.    Claw  tuft  of  Clubiona  obesa  Hentz,  lateral  view,  diagrammatic.   2.  Claw  tuft  of  Aystia  gracilis 
(Hentz),  lateral  view,  diagrammatic.    3.    Generalized   palpal  structure  of  Anyphaena. 


men  of  the  male  is  vibrated  up  and  down 
so  rapidly  that  only  a  blur  is  visible 
(Thompson,  G.  H.,  and  E.  R.  Skinner, 
Courtship  in  Spiders,  Oxford  Scientific 
Films).  Although  the  mating  behavior  of 
very  few  species  in  either  group  has  been 
studied  in  detail,  the  vast  difference  be- 
tween anyphaenid  courtship  and  the  rather 
sluggish  courtship  behavior  of  Clubiona 
and  related  genera  wovild  seem  to  provide 
additional  evidence  for  separating  the  two 
groups  (Platnick,  1971). 

Evidence  that  is  probably  just  as  impor- 
tant as  the  tracheae  for  considering  Any- 
phaenidae  a  distinct  group  is  provided  by 
the  claw  tufts.  Clubionids  have  claw  tufts 
that  are  composed  of  numerous  straight 
simple  setae  densely  clumped  together 
( Text-fig.  1 ) .  Anyphaenid  claw  tufts,  how- 
ever, are  composed  of  two  rows  of  large, 
lamelliform  setae  that  are  greatly  expanded 
at  their  distal  ends  (Text-fig.  2).  All  the 
anyphaenids  examined  have  these  peculiar, 


easily  recognizable  claw  tufts,  but  so  far 
as  known,  no  clubionids  do,  though  some 
phruroliths  have  superficially  similar  claw 
tufts. 

For  these  reasons,  Anyphaenidae  is  here 
considered  a  distinct  family  not  very 
closely  related  to  any  of  the  groups  cur- 
rently included  in  the  Clubionidae. 

RELATIONSHIPS  OF  THE  FAMILY 
ANYPHAENIDAE 

In  addition  to  the  clubionids,  the  any- 
phaenids have  been  associated  with  three 
other  families  of  spiders:  Amaurobiidae, 
Argyronetidae  and  Amaurobioididae.  Leh- 
tinen  (1967)  placed  the  anyphaenids  in 
his  branch  Amaurobiides  and  stated  that 
they  are  probably  derived  from  Amauro- 
biidae: Macrobuninae  and  therefore  lack 
ecribellate,  two-clawed  relatives.  Forster 
( 1970 )  agreed  with  the  placement  of  Any- 
phaenidae  in   Amaurobiides    (and   specifi- 


Spider  Family  Axypiiaenidae  •  Plalnick 


209 


cally  included  the  family  in  his  supeifainily 
Dictynoidca)  but  cited  the  families  Ar- 
gyionetidae  and  Amaurobioididae  as  close 
relatives.  Representatives  of  all  three  fami- 
lies were  examined  to  determine  the  degree 
of  their  relationship,  if  any,  to  the  Any- 
phaenidae. 

Lehtinen  gave  no  evidence  for  his  state- 
ment that  the  anyphaenids  are  probably 
deri\'atives  of  Amaurobiidae:  Macrobuni- 
nae,  presumably  because  there  seems  to  be 
none.  An  examination  of  specimens  of 
one  genus  in  this  subfamily,  Arctohius 
Lehtinen,  indicates  that  it  would  be  diffi- 
cult to  find  araneomorph  spiders  less  likely 
to  have  given  rise  to  the  anyphaenids.  The 
species  of  Arctohius  are  cribellate,  three- 
clawed  spiders  that  lack  claw  tufts  and 
possess  an  unelaborated  tracheal  system. 
Further,  the  genitalia  show  no  similarities 
to  those  of  anyphaenids. 

Likewise,  Forster  gave  no  evidence  for 
associating  the  family  Argyronetidae  with 
tlie  an\q3haenids;  his  decision  to  do  so  was 
based,  I  believe,  on  the  similarities  in  the 
tracheal  systems  of  the  two  groups.  The 
elaboration  of  the  tracheal  system  in  Ar- 
gyroneta,  however,  is  probably  associated 
with  their  invasion  of  an  aquatic  habitat 
and  the  resultant  demands  on  the  respira- 
tory system.  All  the  other  characters,  in- 
cluding the  three  claws,  lack  of  claw  tufts 
and  the  characteristic  pattern  of  ti'icho- 
bothria  distribution,  indicate  that  Argt/ro- 
neta  is,  as  it  is  usually  regarded,  a  close 
relative  (if  not  actually  a  member)  of  the 
family  Agelenidae. 

The  family  Amaurobioididae  was  cre- 
ated by  Hickman  ( 1949 )  for  the  single 
genus  Ammirohioides  O.  P. -Cambridge, 
which  has  at  various  times  been  included 
in  the  families  Drassidae  ( =  Gnaphosi- 
dae),  Ctenidac,  Clubionidae  and  Miturgi- 
dae.  The  genus  is  known  from  New  Zea- 
land, Tasmania,  southern  Chile  and  South 
Africa.  The  spiders  live  in  rock  crevices 
in  the  tidal  zone,  where  they  build  tubular 
silk  retrtnits  and  are  regularly  submerged 
at  high  tide  (Lamoral,  1968). 


Specimens  of  this  rare  genus  provided  by 
R.  R.  Forster  revealed  not  only  a  typically 
anyphac>nid-]ike  tracheal  system,  but  also 
the  lamelliform  claw  tufts  so  characteristic 
of  anyphaenids.  Further,  the  genitalia  are 
close  to  those  of  the  anyphaenid  genus 
Oxysouui,  and  the  body  form  is  similar  to 
that  of  several  species  of  anyphaenids 
known  from  Chile,  Peru,  and  Argentina. 
For  these  reasons,  the  family  Amauro- 
bioididae is  newly  synonymized  with 
Anyphaenidae  in  the  taxonomic  section  of 
this  paper. 

Thus  the  problem  of  the  correct  macro- 
taxonomic  placement  of  Anyphaenidae  has 
been  clarified  but  not  solved  by  this  study 
of  the  groups  with  which  the  family  has 
been  associated  in  the  past.  Futiu-o  work 
should  start  with  an  examination  of  the 
family  Miturgidae  (as  construed  by  Leht- 
inen ) . 

Although  it  was  necessary  to  limit  the 
scope  of  the  detailed  revision  to  the  man- 
ageable number  of  species  occurring  north 
of  Mexico,  all  available  specimens  from 
other  areas  were  examined  to  gain  an 
overview  of  the  family.  Preliminary  im- 
pressions indicate  that  the  family  probably 
originated  in  the  southern  half  of  South 
America  with  subsequent  radiations  north- 
ward. As  indicated  by  the  ability  of 
Amaurohioides  to  withstand  prolonged  sub- 
mersion, it  is  likely  that  early  anyphaenids 
were  able  to  survive  hydrochore  dispersal 
by  rafting,  etc.,  across  considerable  ex- 
panses of  water. 

GENERIC  PROBLEMS  IN  THE 
ANYPHAENIDAE 

The  generic  taxonomy  of  anyphaenids 
is  currently  chaotic.  Every  author  who  has 
worked  with  the  group,  including  Petrunke- 
vitch  (1930),  Bryant  (1931)  and  Chicker- 
ing  (1937),  has  expres.sed  frustration  at 
the  confusion  and  ambiguity  in  the  use  of 
many  of  the  most  common  generic  names. 
One  of  the  principal  causes  of  this  con- 
fusion is  the  interesting  e\olutionary  pat- 
tern   encountered   time    and    again    witliin 


210         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


this  family:  species  tend  to  occur  in  groups 
that  are  remarkably  homogeneovis  in 
genitalic  structure  but  quite  distinct  from 
other  such  groups.  Often  many  of  the 
species  in  these  groups  are  sympatric,  are 
found  in  a  rather  limited  area  and  are 
clearly  the  result  of  radiation  within  that 
area.  An  excellent  example  of  this  is  the 
occurrence  of  nine  closely  related  species 
of  the  Anyphaena  celer  species  group  in 
the  mountains  of  southeastern  Arizona.  It 
is  tempting  to  consider  each  of  these 
groups  a  genus,  as  unambiguous  key  char- 
acters are  then  available  to  distinguish 
genera.  Such  an  approach  would  at  least 
double  the  number  of  genera  found  in  the 
United  States,  and,  if  applied  to  the  Central 
and  South  American  fauna,  would  neces- 
sitate the  creation  of  a  vast  number  of 
new  genera.  If,  instead,  characters  refer- 
ring to  the  general  body  form  are  used,  a 
more  workable  classification  in  terms  of 
both  number  and  size  of  genera  results. 
Unfortunately,  this  makes  the  unambiguous 
definition  of  genera  much  more  difficult 
and  makes  keys  to  genera  awkward  and 
cumbersome.  With  either  approach,  how- 
ever, reliable  genera  composed  of  mo- 
nophyletic  groups  of  species  can  be  estab- 
lished. 

The  second  approach  to  anyphaenid  clas- 
sification has  been  taken  by  the  majority 
of  former  authors,  and  is  continued  in  this 
work.  Thus  the  European  genus  Any- 
phaena is  used  for  the  bulk  of  the  any- 
phaenids  occurring  in  the  United  States, 
even  though  only  one  of  our  species,  Any- 
phaena aperta,  is  actually  a  close  relative 
of  the  European  Anyphaena  accentuata, 
type  species  of  the  genvis.  Nonetheless, 
all  the  species  here  included  in  Anyphaena 
share  a  basic  body  form.  The  neotropical 
genus  Wulfila  is  used  for  all  the  pale, 
long-legged  species,  even  though  they  are 
genitalically  quite  diverse;  the  other  genera 
used  here  are  similarly  construed.  Although 
this  system  is  not  wholly  satisfactory,  it 
seems  decidedly  better  than  creating  a 
host  of  new  generic  names  that  are  likely 


to  fall  into  synonymy  when  a  detailed  ge- 
neric revision  of  the  group  as  a  whole  can 
be  carried  out. 

METHODS 

Tracheae  were  examined  by  dissecting 
away  the  dorsal  cuticle  of  the  abdomen 
and  boiling  the  spider  in  ten  percent 
sodium  hydroxide  for  ten  minutes.  By  this 
method,  all  the  soft  structures  in  the 
abdomen  are  digested  away,  leaving  the 
tracheae  intact. 

Types  of  the  new  species  are  being 
deposited  in  the  American  Museum  of 
Natural  History,  New  York  City,  and  the 
Museum  of  Comparative  Zoology,  Harvard 
University.  Type  depositories  are  abbrevi- 
ated as  follows:  AMNH — American  Mu- 
seum of  Natural  History,  BMNH — British 
Museum,  Natural  History,  MCZ — Museum 
of  Comparative  Zoology. 

Measurements  and  drawings  were  made 
with  a  standard  ocular  grid.  Measurements 
of  gross  morphological  featiu-es  are  ac- 
curate to  ~  0.04  mm;  measurements  of 
ocular  featiu'es  are  accurate  to  ~  0.01  mm. 
Rather  than  selecting  a  small  number  of 
measurements  and  providing  means  and 
standard  deviations  for  these  on  the  basis 
of  a  small  series  of  specimens,  one  male 
and  one  female  of  each  species  were 
measured  in  detail.  As  only  one  of  the 
species  included  here  shows  any  significant 
variation  in  size,  this  procedure  was  deemed 
more  informative.  Actual  measurements 
are  given  rather  than  ratios  since  in  many 
cases  (e.g.,  Anyphaena  catalina  and  A. 
arhida)  closely  related  species  differ  sig- 
nificantly in  size  but  not  in  their  relative 
proportions.  Most  of  the  measurements 
taken  are  self-explanatory,  though  a  few 
need  furdier  comment.  Cephahc  width 
refers  to  the  width  of  the  carapace  at  a 
point  just  behind  the  posterior  median 
eyes,  and  thus  provides  an  indication  of 
the  degree  to  which  the  carapace  is  nar- 
rowed in  front. 

The  difficult  problem  of  accurately  de- 
scribing   die    eye    relationships    has    been 


Spider  Family  Anyphaenidae  •  Platnick        211 


solved  by  providing  a  set  of  measurements 
from  wliich  it  is  possible  to  reconstrnct, 
using  grapli  paper,  the  exact  eye  arrange- 
ment. Diameters  are  given  using  the  con- 
ventional abbreviations  (AME  =  anterior 
median  eye,  ALE  =  anterior  lateral  eye, 
PME  =  posterior  median  eye,  PLE  = 
posterior  lateral  eye).  The  length  of  each 
eye  row  is  measured  from  the  lateral  edge 
of  one  lateral  eye  to  the  lateral  edge  of 
tlie  other  lateral  eye.  Curvature  of  the  eye 
rows  is  described  as  viewed  frontally,  not 
dorsally.  This  was  accomplished  by  posi- 
tioning the  spider  in  sand,  a  technique 
found  most  useful  for  making  all  the 
measurements.  The  dimensions  of  the 
median  ocular  quadrangle  (MOQ)  are 
given,  as  well  as  the  distances  between 
each  of  the  eyes.  The  latter  measurements 
extend  between  the  edges  of  the  lenses  of 
the  eyes  under  consideration  (not  just 
between  the  dark  circles  surrounding  each 
eye). 

The  relative  length  and  thickness  of  each 
leg  is  indicated  by  the  tibial  length  index — 
the  tibial  width  divided  by  the  tibial 
length,  with  the  result  multiplied  by  100 
to  obtain  a  whole  number.  All  tibial 
measurements  were  taken  from  a  dorsal 
view  and  refer  to  the  maximum  lengths 
and  widths.  The  lower  the  tibial  index, 
the  longer  and  thinner  the  leg;  conversely, 
the  higher  the  index,  the  shorter  and  thicker 
the  leg.  In  practice  the  index  varies  from 
around  3  to  35. 

Ventral  spination  of  the  leg  segments  is 
indicated  by  the  standard  formula  in  which 
the  number  of  spines  on  the  proximal, 
median  and  distal  thirds  of  the  leg  segment 
are  given.  Only  ventral  spines,  not  lateral 
ones,  are  included,  and  any  even  number 
in  the  formula  may  be  taken  to  represent 
a  pair  of  spines.  Unless  the  last  number  is 
followed  by  an  asterisk,  the  last  pair  of 
spines  is  terminally  located.  Thus,  for 
example,  the  formula  2-2-2*  indicates  that 
the  segment  bears  three  pairs  of  ventral 
spines,  the  last  pair  of  which  is  not  termi- 
nally located.    The  term  "spine"  is  used  in 


its  conventional  arachnological  sense  and 
rc'fers  to  the  moxable  macrosetae  found 
on  the  legs.  Similarly,  the  term  "clypeus" 
is  used  to  refer  to  the  area  between  the 
anterior  eye  row  and  the  anterior  edge  of 
tlie  carapace  and  not  to  the  small  sclerite 
folded  under  the  carapace.  Since  neither 
usage  of  tht>  term  reflects  certain  knowledge 
of  homology  with  the  insect  clypeus,  the 
old  and  established  usage  should  be  main- 
tained. 

Scale  lines  for  the  drawings  always  equal 
0.1  mm.  Each  scale  line  applies  to  all 
consecutively  numbered  drawings  imtil  a 
new  scale  line  appears.  Exceptions  are 
noted  in  the  captions. 

TAXONOMY 
Anyphaenidae 

Anyphaenidae  Bertkau,  1878,  Arch.  Naturg.,  44: 
358,  379.  Tvpe  genus  Anyphaena  Sundevall, 
1833. 

Amaurobioididae  Hickman,  1949,  Pap.  Proc.  Roy. 
Soc.  Tasmania,  1948:  31.  Tvpe  genus  A77iauro- 
hioides  O.P.-Cambridge,  1883.  NEW  SYN- 
ONYMY. 

Diagnosis.  The  combination  of  the  ad- 
vanced tracheal  spiracle  and  the  lamelli- 
form  claw  tufts  will  serve  to  distinguish  the 
anyphaenids    from    all    other   families. 

Description.  Chelicerae  diaxial,  not  fused 
together  at  base.  Labium  free.  Without 
cribellum  or  calamistrum.  With  one  pair 
of  book  lungs  and  a  tracheal  spiracle  lo- 
cated considerably  anterior  to  the  spin- 
nerets, most  often  midway  between  spin- 
nerets and  epigastric  furrow,  sometimes 
closer  to  one  or  the  other.  Eight  eyes  in 
two  rows.  Six  spinnerets,  anterior  spin- 
nerets approximate,  colulus  represented 
only  by  hairs,  anal  tubercle  unmodified. 
Legs  prograde,  metatarsi  and  tarsi  I  and 
II  scopulate,  tarsi  with  two  toothed  claws 
and  claw  tufts  composed  of  lamelliform 
setae. 

Key  to  Genera 
IN  America  north  of  Mexico 

la.  Tracheal  spiracle   luucli   closer  to  epigastric 
furrow   tlian   to   spinnerets   ..Aysha 


212         Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  4 


lb.  Tracheal  spiracle  roughly  midway  between 
epigastric   furrow   and   spinnerets   2 

2a.  Legs  very  long  and  thin.  Leg  I  greatly 
elongated,  tibial  index  (width/length  X 
100)   usually  5  or  less  Wulfila 

2b.  Legs  normal,  tibial  index  of  leg  I  usually 
8    or    more    3 

3a.  Chelicerae  with  2  retromarginal  teeth  

Oxysoma 

3b.  Chelicerae  with  4-9  retromarginal  denticles 
4 

4a.  Carapace  usually  with  two  dark  paramedian 
longitudinal  bands;  chelicerae  not  produced 
forward;  femora  not  much  darker  than 
other  leg  segments  Anijphaena 

4b.  Carapace  without  dark  paramedian  longi- 
tudinal bands;  either  chelicerae  produced 
forward  or  femora  much  darker  than  other 
leg   segments Teudis 

Anyphaena  Sundevall 

Amjphaena  Sundevall,  1833,  Conspectus  Arachn., 
28.  Type  species  by  monotypy  Aranea  ac- 
centuata  Walckenaer,   1802. 

Diagnosis.  The  combination  of  the  fol- 
lowing characters  will  serve  to  distinguish 
the  genus  in  America  north  of  Mexico: 
trachael  spiracle  roughly  midway  between 
epigastric  furrow  and  spinnerets,  leg  I  not 
greatly  elongated,  chelicerae  with  4-9 
retromarginal  denticles  and  not  produced 
forward,  femora  not  much  darker  than 
other  leg  segments.  The  carapace  usually 
has  two  dark  paramedian  longitudinal 
bands.  The  genus  is  used  here  in  a  very 
broad  sense;  this  prevents  simple  diagnosis, 
and  makes  detailed  descriptions  of  each 
species  group  more  meaningful  than  a 
description  of  the  whole  genus. 

Uncertain  names.  Types  of  the  follow- 
ing species  were  unavailable  and  are  too 
poorly  described  to  permit  identification: 
Cluhiona  agresiis  Hentz,  1847,  type  de- 
stroyed; Chihiona  fallens  Hentz,  1847,  type 
destroyed,  Cluhiomi  suhlurida  Hentz,  1847, 
type  destroyed;  Amjphaena  argentata 
Becker,  1879,  type  lost;  and  Amjphaena 
striata  Becker,  1879,  type  lost.  The  three 
Hentz  Cluhiona  species  were  transferred  to 
Anijphaeiui  by  Marx  (1890),  but  there  is 
little    justification    for    this    in    the    vague 


descriptions.     All    the    above    names    are 
regarded  as   nomina  cluhia. 

Species  groups.  Although  there  seem  to 
be  several  species  groups  of  Amjphaena  in 
the  Neotropic  region,  only  four  occur  north 
of  Mexico.  The  celer  group  is  the  largest; 
it  has  representatives  at  least  as  far  south 
as  Panama  and  probably  contains  over 
thirty  species.  The  pectorosa  and  pacifica 
groups  are  closely  related  and  occur  com- 
monly in  Mexico  as  well  as  the  United 
States;  it  is  difficult  to  place  females  in 
one  group  or  the  other  unless  the  male  is 
also  known;  they  probably  contain  together 
at  least  twenty  species.  The  accentuata 
group  is  predominantly  Palearctic  and  prob- 
ably contains  at  least  five  species. 

Key  to  Species  Groups 

la.  Metatarsi  I  and  II  with  one  pair  of  ventral 
spines  accentuata  group 

lb.  Metatarsi  I  and  II  with  two  pairs  of  ventral 
spines    2 

2a.  Retrolateral  tibial  apophysis  of  males  bifid, 
with  ventral  prong  elongated  ( Figs.  18-20, 
25-32).  Epigynum  of  females  with  a  hood 
(Figs.  21,  23,  33,  36,  37,  39-42)  ._  celer  group 

2b.  Retrolateral  tibial  apophysis  of  males  not 
bifid  or  elongated  (Figs.  55-58,  69-71). 
Epigynum   without    a   hood    (Figs.    66,    67, 

72,  74,  77,  79)  3 

3a.  Eastern   United  States.    Coxae   III   and   IV 

of  males  with  pointed  spins  (Figs.  59-62). 
Female  epigyna  on  broad  sclerotized  plates 
(Figs.  74,  77,  79);  internal  genitalia  lacking 

long  ducts   (Figs.  75,  78,  80)    

pectorosa    group 

3b.  Western  United  States.  Coxae  III  and  IV 
of  males  without  pointed  spurs,  though 
rounded  knobs  may  be  present.  Female 
epig>'na  not  on  broad  sclerotized  plates 
(Figs.  66,  67,  72);  internal  genitalia  with 
long,    sometimes    coiling,    ducts    (Figs.    68, 

73,  76)    _._. pacifica  group 

Anyphaena  celer  Group 

Diagnosis.  Males  of  the  celer  group  may 
be  recognized  by  their  retrolateral  tibial 
apophysis,  which  is  usually  bifid  with  an 
elongated  ventral  prong  (Figs.  18,  20,  26). 
Females  have  a  characteristic  epigynum 
consisting  of  a  hood,  two  sidepieces  and  a 
midpiece    (Figs.   9,   33),   though  the   mid- 


SpiDEii  Family  Axyphakxidae  •  Pkilnick        213 


piece  is  reduced  in  A.  crebrispimi  ;ind  A. 
(lixiana  (Figs.  21,  23). 

Description.  Total  length  3-7  nnn,  with 
males  of  most  species  between  3.3-4.6  mm, 
females  of  most  species  between  4.1-5.9 
mm.  Carapace  longer  than  wide,  narrowed 
in  front  to  less  than  half  its  maximum 
w  idth.  Clypens  height  greater  than  anterior 
median  eye  diameter.  Posterior  median, 
posterior  lateral  and  anterior  lateral  eyes 
subeqiial  in  size,  larger  than  anterior 
medians.  Procurved  posterior  eye  row 
longer  than  recin'\'ed  anterior  row.  Median 
ocular  ({nadrangle  longer  than  wide  in 
front,  wider  than  long  in  back.  Anterior 
median  eyes  separated  by  their  diameter, 
by  their  radius  from  anterior  laterals. 
Posterior  medians  separated  by  their  diam- 
eter, slightly  closer  to  posterior  laterals 
than  to  each  other.  Anterior  laterals 
separated  by  their  radius  from  posterior 
laterals.  Sternum  longer  than  wide,  un- 
modified. Chelicerae  with  4-5  promarginal 
teeth  and  6-9  reti-omarginal  denticles. 
Abdomen  longer  than  wide,  tracheal 
spiracle  midway  between  epigastric  fur- 
row and  base  of  spinnerets.  Leg  formula 
1423.  Metatarsi  I  and  II  with  two  pairs 
of  \'entral  spines.  Males  often  with  femur 
III  thickened  distally,  set  with  stiff  short 
setae  ventrally;  tibia  III  ventral  spines 
thickened,  cone-like;  coxae  set  with  clumps 
of  stiff  short  setae.  Palpus  with  an  elon- 
gated median  apophysis,  retrolateral  teg- 
ular apophysis,  conspicuous  curving  em- 
bolus and  conductor.  Retrolateral  tibial 
apophysis  bifid,  with  dorsal  prong  reduced 
in  some  species.  Epigynum  with  hood, 
two  sidepieces  and  midpiece;  two  simple 
spermathecae. 

Variation.  None  of  the  species  in  this 
group  show  any  significant  individual  or 
geographic  intraspecific  variation  in  struc- 
tin-e,  size  or  coloration. 

Key  to  Species 

la.  Dorsal  and  \entral  prongs  of  retrolateral 
tibial  apophysis  ( RTA )  roughly  equal  in 
length  (Figs.  18,  19);  epigynal  hood  wide, 
more  than  four  times  the  minimum  width 


of  epigynal  sidepiece   ( Figs.  9,   11);  east- 
ern   U.S 2 

II).  Ventral  prong  of  retrolateral  tibial  apophy- 
sis ( RTA )  nuieh  longer  than  dorsal  prong 
(as  in  Figs.  26,  27);  epigynal  hood 
narrow,  less  than  four  times  the  minimum 
width  of  epigynal  sidepiece  ( as  in  Figs. 
33,   36);   western   U.S.   3 

2a.  Dorsal  prong  of  RTA  broad,  with  a  trans- 
lucent ridge  (Fig.  18);  epigynal  hood  a 
thick  oval,  sidepieces  straight  (Fig.  9) 
— - - — _..    celer 

21).  Dorsal  prong  of  RTA  narrow,  without  a 
translucent  ridge  (Fig.  19);  epigynal  hood 
a  thin  oval,  sidepieces  rounded  ( Fig.  1 1 ) 
- - _     inaculata 

3a.  Base  of  RTA  expanded  into  a  broad 
triangle  (Fig.  20);  retrolateral  tegular 
apophysis  prolonged  medially  (Fig.  3); 
epigynal  sidepieces  more  than  three  times 
the  width  of  epigynal  hood  (Fig.  21) 
-    crebrispina 

3b.  Base  of  RTA  not  expanded;  retrolateral 
tegular  apophysis  not  prolonged  medialK ; 
epigynal  sidepieces  less  than  three  times 
the  width  of  epigynal  hood  4 

4a.  Dorsal  prong  of  RTA  bearing  a  sharp  spur 
(Fig.  25);  epigynal  midpiece  greatly  re- 
duced, sidepieces  widely  separated  pos- 
teriorly   (Fig.    23)    dixiana 

4b.  Dorsal  prong  of  RTA  without  a  spur; 
epigynal  midpiece  conspicuous,  sidepieces 
approximate  posteriorly  5 

5a.  Males    '. 6 

5b.   Females     14 

6a.  Dorsal  prong  of  RTA  with  two  triangular 
processes  separated  by  a  concave  notch 
( Fig.   26 )    judicata 

6b.  Dorsal  prong  of  RTA  witliout  triangular 
processes     „. 7 

7a.  Dorsal  prong  of  RTA  with  a  long  recurved 
hook  (Fig.  29)   autumna 

7b.  Dorsal  prong  of  RTA  without  a  long  re- 
curved   hook    8 

8a.  Dorsal  prong  of  RTA  witli  a  basal  hook 
(Figs.   31,    38)    _.-.. 9 

8b.  Dorsal  prong  of  RTA  without  a  basal 
hook    10 

9a.  Conductor  and  retrolateral  tegular  apophy- 
sis recurved    (Fig.   15)    catalina 

9b.  Conductor  and  retrolateral  tegular  apoph- 
ysis  not   recurved    ( Fig.    17 )    arhida 

10a.   Dorsal    prong    of    RTA    a    shaiply    pointed 

spike    (  Fig.   32 )    liespar 

101).   Dorsal     prong     of     RTA     not     a     sharply 

pointed  spike 1 1 

11a.  Fmbolus  with   a  conspicuous  enlargement 

(Figs.  7,  13) .12 

111).   Embolus    without    a    con.spicuous    enlarge- 
ment  (Figs.  21,  33) 13 


214         Bulletin  Museum  of  Comparative  Zoolog,ij,  Vol.   146,  No.   4 


J? 


I       ^J'',  ,^-^  m-p. 


Anyphaeno  maculota  \  q\  —>> 


o- 


I     -* 

J 

\  \A- 

1 

1 

^~«    ', 

>-:-: 

Anyphoena  crebrispino 

/— 

\\v 

Anyphaena  dixiana        j»-  \ 


v^\  \ 


Anyphaena  rita    ' 


V    f 

1       ~T 
1            1 

\h~^ 

'             1 

Anyphaena 

cochise V 

Map  1.     Distributions  of  Anyphaena  arbida,  A.  autumna,  A.  catalina,  A.  celer,  A.  cochise,  A.  crebrispina,  A.  dixi- 
ana, A.  gibboides,  A.  hespar,  A.  judicata,  A.  maculata,  A.  marginalis  and  A.  rita. 


12a.   Dorsal  prong  of  RTA  more  than  half  the 

length  of  ventral  prong   ( Fig.  35 )   

cochise 

12b.   Dorsal   prong   of   RTA   less   than   half   the 

length  of  ventral  prong   (Fig.  28)   

rita 

13a.  Median  apophysis  sharply  pointed;  con- 
ductor short,  bent  (Fig.  14);  Oregon  and 
Utah   gibboides 

13b.  Median  apophysis  rounded;  conductor 
long,  straight  (Fig.  6);  Arizona  and  New 
Mexico  marginalis 

14a.  Epigynal  hood  wider  than  long;  midpiece 
not  wider  than  hood,  without  constric- 
tions; sidepieces  very  wide  (Fig.  40); 
Oregon  and   Utah   gibboides 

14b.  Epigynal  hood  as  long  as  wide  or  midpiece 
wider  than  hood  or  sidepieces  narrow; 
Arizona  and  New  Mexico  15 

15a.  Epigynal  midpiece  a  very  broad  triangle 
( Fig.    37 )    rita 

15b.  Epigynal  midpiece  othenvise  16 

16a.  Spermathecae  much  further  apart  pos- 
teriorly than  anteriorly   ( Fig.  49 )   _-,  hespar 

16b.  Spemiathecae  as  far  apart  anteriorly  as 
posteriorly     17 

17a.  Epigynal  hood  much  wider  than  long  ( Fig. 
39 )    _  autumna 

17b.  Epigynal  hood  as  long  as  wide  —18 


18a.  Epigynal    midpiece    less    than    twice    the 

length  of  epigynal  hood  (Fig.  41)  

catalina 

18b.  Epigynal  midpiece  more  than  twice  the 
length   of  epigynal  hood   ...19 

19a.  Epigynal  midpiece  a  short  triangle  (Fig. 
33 )    judicata 

19b.  Epigynal  midpiece  an  elongate  triangle 
(Fig.    36)    marginalis 

Anyphaena  celer  (Hentz) 
Map  1;  Figures  1,  9,  10,  18 

Chibiona  celer  Hentz,  1847,  J.  Boston  Soc.  Natur. 
Hist.,  5:  452,  pi.  23,  fig.  20  (  9  ).  Male  holo- 
type,  female  allotype  from  Alabama  and  North 
Carolina  in  the  Boston  Soc.  Natur.  Hist. 
(Boston  Museum  of  Science),  destroyed  by 
beetles. 

Anyphaena  incerta  Keyserling,  1887,  Verb.  zool. 
hot.  Ces.  Wien,  .37:  452,  pi.  6,  fig.  22  (  $  ). 
Female  holotype  from  Cambridge,  Massachu- 
setts, in  MCZ,  examined.  Emerton,  1890,  Trans. 
Connecticut  Acad.  Sci.,  8:  186,  pi.  6,  figs. 
2-2d,  $,9. 

Anyphaena  celer,  Simon,  1897,  Hist.  Natur. 
Araign.,  2:  96.  Bryant,  1931,  Psyche,  38:  111, 
pi.  6,  fig.  9,  pi.  8,  figs.  25,  28,  $,  9.  Chickering, 
1939,   Pap.   Michigan   Acad.    Sci.,   24:    51,   figs. 


Spider  Family  Axyphakxidae  •  PJaliiick        215 


Plate  1 
Figures  1-8.    Left  palpi,  ventral  view.    Figures  9.  11.    Epigyna,  ventral  view.    Figures  10,  12.    Internal  genitalia, 
dorsal  view.     1,9,10.   /Anyphaena  ce/er  (Hentz).     2,11,12.    Anyphaena  maculata  (Banks).     3.    Anyphaena  crebri- 
sp'ma  Chamberlin.    4.  Anyphaena  dixiana  (Chamberlin  and  Woodbury).    5.    Anyphaena  judicata  O.  P. -Cambridge. 
6.   Anyphaena  marginalis  (Banks).    7.   Anyphaena  rita  new  species.    8.   Anyphaena  autumna  new  species. 


216         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


t>' 


1-4,  $,  9.  Comstock,  1940,  Spicier  Book, 
rev.  ed.,  p.  577,  figs.  634-6.35,  $,9.  Kaston, 
1948,  Bull.  Connecticut  Geol.  Natur.  Hist. 
Surv.,  70:  407,  figs.  1471-1476,   $,9.  Roewer, 

1954,  Katalog    der    Araneae,    2:528.      Bonnet, 

1955,  Bibliographia   Araneorum,   2:    343. 
Gatjenna  celer,   Comstock,    1912,   Spider   Book,   p. 

563,  figs.  634-635,  $,9. 

Dia<ino.sis.  Anyphaena  celer  is  most 
closely  related  to  A.  maculata.  Males  of 
both  species  have  dorsal  and  venti-al  RTA 
prongs  roughly  equal  in  length,  but  A.  celer 
males  may  be  distinguished  by  the  trans- 
lucent ridge  on  their  dorsal  prong  (Fig. 
18).  Females  may  be  separated  by  the 
straight  epigynal  sidepieces  and  widely 
oval  epigynal  hood  of  A.  celer  (Fig.  9). 

Male  (Jackson  Co.,  Illinois).  Total 
length  4.54  mm.  Carapace  2.12  mm  long 
1.58  mm  wide,  cephalic  width  0.83  mm 
clypeus  height  0.07  mm,  pale  yellow  with 
thin  dark  broken  border  and  two  dark 
paramedian  longitudinal  bands.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.12, 
PME  0.11,  PLE  0.12;  anterior  eye  row 
0.44  mm  long,  slightly  recurved;  pos- 
terior eye  row  0.60  mm  long,  procurved; 
MOQ  length  0.24  mm,  front  width  0.19 
mm,  back  width  0.31  mm;  eye  interdis- 
tances  (mm):  AME-AME  0.06,  AME- 
ALE  0.03,  PME-PME  0.10,  PME-PLE 
0.10,  ALE-PLE  0.05. 

Sternum  1.06  mm  long,  0.94  mm  wide, 
pale  yellow  with  dark  markings  opposite 
coxae,  translucent  border  and  darkened 
extensions  to  coxae.  Chelicerae  0.79  mm 
long  with  4  promarginal  teeth  and  8  retro- 
marginal  denticles,  pale  yellow  with  boss 
outlined  in  gray.  Labium  and  endites  pale 
yellow,  darkest  proximally;  endites  not 
invaginated. 

Abdomen  2.30  mm  long,  1.62  mm  wide, 
pale  white  with  transverse  rows  of  dark 
markings;  venter  with  scattered  dark  mark- 
ings. Epigastric  furrow  0.85  mm  from 
tracheal  spiracle,  spiracle  0.88  mm  from 
base  of  spinnerets. 

Legs  pale  yellow  with  scattered  dark 
markings.  Tibial  lengths  (mm)  and 
indices:  I  1.98,  12;  II  1.82,  14;  III  1.17,  21; 


IV  1.73,  16.  Ventral  spination:  tibiae  I-IV 
2-2-2;  metatarsi  I,  II  2-2-0,  III  2-0-2,  IV 
2-2-2.  Femur  III  thickened  distally  with 
clump  of  short  thick  setae  ventrally.  Tibia 
III  ventral  spines  1,  2  on  retrolateral  side 
thickened,  cone-like.  Coxae  III,  IV  pro- 
lateral  ventral  surface  with  clump  of  short 
thick  setae. 

Palpus  as  in  Figures  1,  18. 

Female  (Wayne  Co.,  Ohio).  Coloration 
as  in  male.  Total  length  5.87  mm.  Carapace 
2.07  mm  long,  1.39  mm  wide,  cephalic 
width  0.86  mm,  clypeus  height  0.05  mm. 
Eyes:  diameters  (mm):  AME  0.05,  ALE 
0.10,  PME  0.10,  PLE  0.10;  anterior  eye  row 
0.42  mm  long,  recurved;  posterior  eye  row 
0.58  mm  long,  procurved;  MOQ  length 
0.30  mm,  front  width  0.19  mm,  back  width 
0.32  mm;  eye  interdistances  (mm):  AME- 
AxME  0.07,  AME-ALE  0.05,  PME-PME 
0.12,  PME-PLE  0.07,  ALE-PLE  0.08. 

Sternum  0.99  mm  long,  0.88  mm  wide. 
Chelicerae  0.76  mm  long  with  teeth  as  in 
male. 

Abdomen  4.10  mm  long,  2.13  mm  wide. 
Epigastric  furrow  1.57  mm  from  tracheal 
spiracle,  spiracle  1.37  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.58,  15;  II  1.42,  16;  III 
1.01,  24;  IV  1.60,  15.  Ventral  spination: 
tibiae  I,  II  2-2-2,  III,  IV  1-1-2;  meta- 
tarsi I,  II  2-2-0,  III  2-0-2,  IV  2-2-2. 

Epigynum  as  in  Figure  9,  internal  geni- 
talia as  in  Figure  10. 

Natural  history.  Mature  males  have  been 
taken  every  month  except  June,  mature 
females  year-round.  Specimens  have  been 
taken  in  houses,  deciduous  forests,  on 
leaves,  flowers,  ti-eesides,  in  pitfalls  and 
footprints  in  snow. 

Distribution.  Eastern  United  States  from 
southern  New  England  west  to  Wisconsin, 
south  to  Florida  and  Texas   (Map  1). 

Anyphaena  maculata  (Banks) 
Map  1;  Figures  2,  11,  12,  19 

Caijcnna  maculata  Banks,  1896,  Trans.  Amer. 
Ent.  Soc,  23:  64.  Male  holotype  from  Wash- 
ington,   D.C.,   in    MCZ,   examined.     Bishop   and 


Spider  Family  Anyphaenidak  •  Plat  nick        217 


N^^ 


Figures  13-17.    Left  palpi,  ventral  view, 
ventral    view.      Figures    22,    24.     Interna 
Anyphaena  gibboides  new  species.     15. 
17.    Anyphaena  arbida  new  species.     18 
22.   Anyphaena  crebrispina  Channberlin. 


Plate  2 
Figures  18-20.    Left  palpi,  retrolateral  view.    Figures  21,  23.    Epigyna, 

genitalia,    dorsal   view.     13.    Anyphaena   cochise   new   species.     14. 
Anyphaena  catalina  new  species.     16.   Anyphaena  hespar  new  species. 

Anyphaena  celer  (Hentz).     19.    Anyphaena  maculata  (Banks).     20.  21, 
23,  24.    Anyphaena  dixlana  (Chamberlin  and  Woodbury). 


218         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


Plate  3 
Figures  25-32.  Left  palpi,  retrolateral  view.  Figure  33.  Epigynum,  ventral  view.  Figure  34.  Internal  genitalia, 
dorsal  view.  25.  Anyphaena  dixiana  (Chamberlin  and  Woodbury).  26,  33,  34.  Anyphaena  judicata  O.  P. -Cam- 
bridge. 27.  Anyphaena  marginalis  (Banks).  28.  Anyphaena  rita  new  species.  29.  Anyphaena  autumna  new 
species.  30.  Anyphaena  gibboides  new  species.  31.  Anyphaena  catalina  new  species.  32.  Anyphaena  hespar 
new  species. 


Spider  Family  Anyphaenidae  •  Platnick        219 


Crosby,  1926,  T.  Elislia  Mitclioll  Sci.  Soc,  41: 
189,  pi.  24,  figs.  37,  38,  $,  9. 
Amjphacna  nuiciihita,  Simon,  1897,  Hist.  Natiir. 
Araign.,  2:  96.  Brvant,  1931,  Psyche,  38:  111, 
pi.  6,  fig.  8,  pi.  8,  fig.  31,  $,  9.  Kaston,  1948, 
Bull.  Connecticut  Geol.  Natur.  Hist.  Surv., 
70:  409,  figs.  1457-1458,  $,9.  Roewer,  1954, 
Katalog  der  Araneae,  2:  529.  Bonnet,  1955, 
Bibliographia  Araneoruni,  2:  345. 

Diagnosis.  Anyphoena  inaculata  is  most 
closely  related  to  A.  celer.  Males  may  be 
distinguished  by  the  short  dorsal  prong  of 
the  RTA,  which  lacks  a  translucent  ridge 
(Figure  19);  females  by  their  rounded 
epigynal  sidepieces  and  narrowly  oval 
epigynal    hood    (Figure    11). 

Male  (Durham  Co.,  North  Carolina). 
Coloration  as  in  Amjphaena  celer.  Total 
length  3.74  mm.  Carapace  2.09  mm  long, 
1.54  mm  wide,  cephalic  width  0.77  mm, 
clvpeus  height  0.08  mm.  Eyes:  diameters 
(mm):  AME  0.07,  ALE  0.11,  PME  0.10, 
PLE  0.10;  anterior  eye  row  0.44  mm  long, 
recurved;  posterior  eye  row  0.58  mm  long, 
procurved;  MOQ  length  0.23  mm,  front 
width  0.20  mm,  back  width  0.31  mm;  eye 
interdistances  (mm):  AME-AME  0.05, 
AME-ALE  0.03,  PME-PME  0.10,  PME- 
PLE  0.10,  ALE-PLE  0.04. 

Sternum  1.08  mm  long,  0.79  mm  wide. 
Chelicerae  0.63  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  2.02  mm  long,  1.08  mm  wide. 
Epigastric  furrow  0.31  mm  from  tracheal 
spiracle,  spiracle  0.41  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.00, 
11;  II  1.75,  13;  III  1.12,  24;  IV  1.82,  15. 
Ventral  spination:  tibiae  I,  II  2-2-2*,  III, 
IV  2-2-2;  metatarsi  I  2-1-0,  II  2-2-0,  III 
2-0-2,  IV  2-2-2.  Modifications  of  third 
leg  as  in  A.  celer. 

Palpus  as  in  Figures  2,  19. 

Female  (Pope  Co.,  Illinois).  Coloration 
as  in  male  of  A.  celer. 

Total  length  4.68  mm.  Carapace  2.07 
mm  long,  1.60  mm  wide,  cephalic  width 
0.97  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.08,  ALE  0.10, 
PME  0.10,  PLE  0.11;  anterior  eye  row  0.48 


mm  long,  recurved;  posterior  eye  row  0.63 
mm  long,  procur\'ed;  MOQ  length  0.30 
mm,  front  width  0.22  mm,  back  width  0.33 
nun;  eye  interdistances  (mm):  AME- 
AME  0.07,  AME-ALE  0.04,  PME-PME 
0.14,  PME-PLE  0.11,  ALE-PLE  0.05. 

Sternum  1.15  mm  long  and  0.95  mm 
wide.  Chelicerae  0.71  mm  long  with  teeth 
as  in  male. 

Abdomen  3.02  mm  long,  2.11  nun  wide. 
Epigastric  furrow  0.85  mm  from  tracheal 
spiracle,  spiracle  0.90  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (nun) 
and  indices:  I  1.42,  18;  II  1.48,  18;  III  0.99, 
25;  IV  1.58,  16.  Ventral  spination:  tibiae 
I,  II  2-2-2',  III  1-1-2,  IV  2-1-2;  metatarsi 
I,  II  2-2-0,  III  2-0-2,  IV  2-2-2. 

Epigynum  as  in  Figure  11,  internal  geni- 
talia as  in  Figure  12. 

Natural  history.  Mature  males  have  been 
taken  from  late  September  through  early 
February,  mature  females  from  mid-Octo- 
ber through  mid- April.  Specimens  have 
been  taken  from  Spanish  moss,  by  sweep- 
ing in  bottomland  pine  and  hardwood 
forests,  by  sifting  leaves  and  by  Malaise 
trap. 

Distribution.  Mid-eastern  states  from 
Long  Lsland  south  to  North  Carolina,  wost 
to  southern  Illinois,  eastern  Missouri  and 
northern  Alabama  ( Map  1 ) . 

Anyphaena  crebrispina  Chamberlin 
Map  1;  Figures  3,  20,  21,  22 

Aiufpliacna  crebrispina  Chamberlin,  1919,  Pomona 
Coll.  J.  Ent.  Zool.,  12:  10,  pi.  4,  fig.  4  {$). 
Male  holotype  from  Clareniont,  California,  in 
MCZ,  examined.  Bryant,  1931,  Psyche.  38:  113, 
pi.  6,  fig.  11,  $.  Roewer,  1954,  Katalog  der 
Araneae,  2:  528.  Bonnet,  1955,  Bibliographia 
Araneoruni,  2:  343. 

Amjphaena  zina  Chamberlin,  1919,  Pomona  Coll. 
J.  Ent.  Zool.,  12:11,  pi.  4,  fig.  5(9).  Female 
holotype  from  Clareniont,  California,  in  MCZ, 
examined.  Roewer,  1954,  Katalog  der  Araneae, 
2:  530.  Bonnet,  1955,  Bibliographia  Araneorum, 
2:  349.  NEW  SYNONYMY. 

Diagnosis.  Anyphaena  crebrispina  is  the 
most  aberrant  member  of  the  celer  group, 
but  is  most  closely  related  to  A.  dixiatm. 


220         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


Plate  4 
Figures  35,  38.  Left  palpi,  retrolateral  view.  Figures  36,  37,  39-42.  Epigyna,  ventral  view.  Figures  43-46,  48, 
49.  Internal  genitalia,  dorsal  view.  Figure  47.  Anyphaenid  tracheae,  diagrammatic.  Figure  50.  Clubionid 
tracheae,  diagrammatic.  35.  Anyphaena  cochise  new  species.  36,  43.  Anyphaena  marginalis  (Banks).  37,  44. 
Anyphaena  rita  new  species.  38.  Anyphaena  arbida  new  species.  39,  45.  Anyphaena  autumna  new  species. 
40,46.  Anyphaena  gibboides  new  species.  41,48.  /Anyp/7aeA?a  ca?a//na  new  species.  42,49.  Anyphaena  hespar 
new  species. 


Spidkh  I-'amily  Anyphaexidak  •  I'latnick        221 


Males  of  A.  cre])ris-}nna  may  be  readily  dis- 
tinguished by  the  greatly  expanded  base 
of  the  RTA  (Fig.  20).  If' this  speeies  were 
known  solely  from  the  female,  it  would  be 
impossible  to  place  it  in  the  celer  group: 
the  epigynum,  with  its  greatly  expanded 
sidepieces  and  its  hick  of  an  c>xternally  visi- 
ble midpiece,  is  totally  unlike  that  of  any 
other  species  in  this  group  ( Fig.  21 ) . 

Male  (Los  Angeles  Co.,  California). 
Coloration  as  in  AnypJiaena  celer.  Total 
length  4.61  mm.  Carapace  2.00  mm  long, 
1.57  mm  wide,  cephalic  width  0.74  mm, 
clypeus  height  0.10  mm.  Eyes:  diameters 
(mm):  AME  0.07,  ALE  0.10,  PME  0.10, 
PLE  0.10;  anterior  eye  row  0.43  mm  long, 
recurved;  posterior  eye  row  0.56  mm  long, 
prociuved;  MOQ  length  0.25  mm,  front 
width  0.18  mm,  back  width  0.29  mm;  eye 
interdistances  (mm):  AME-AME  0.04, 
AME-ALE  0.02,  PME-PME  0.09,  PME- 
PLE  0.09,  ALE-PLE  0.04. 

Sternum  1.10  mm  long,  0.S8  mm  wide. 
Chelicerae  0.55  mm  long  with  5  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  2.65  mm  long,  1.58  mm  wide. 
Epigastric  furrow  0.79  mm  from  tracheal 
spiracle,  spiracle  0.68  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  L69, 
13;  II  1.51,  15;  III  1.06,  22;  IV  1.69,  14. 
Ventral  spination:  tibiae  I  2-2-2*,  II  1-2- 
2\  III  1-2-2,  IV  2-2-2;  metatarsi  I,  II  2- 
2-0,  III  2-0-2,  IV  2-2-2.  Modifications  of 
third  leg  as  in  A.  celer. 

Palpus  as  in  Figures  3,  20. 

Female  (Los  Angeles  Co.,  California). 
Coloration  as  in  male  of  A.  celer.  Total 
length  4.39  mm.  Carapace  1.85  mm  long, 
1.37  mm  wide,  cephalic  width  0.77  mm, 
clypeus  height  0.08  mm.  Eyes:  diameters 
(mm):  AME  0.07,  ALE  0.09,  PME  0.09, 
PLE  0.09;  anterior  eye  row  0.41  mm  long, 
recurved;  posterior  eye  row  0.56  mm  long, 
procurved;  MOQ  length  0.23  mm,  front 
width  0.18  mm,  back  width  0.28  mm;  eye 
interdistances  (mm):  AME-AME  0.04, 
AME-ALE  0.02,  PME-PME  0.10,  PME- 
PLE  0.08,  ALE-PLE  0.04. 


Sternum  1.08  mm  long,  0.86  mm  wide. 
Chelieerac>  0.64  mm  long  with  4  promar- 
ginal  teeth  and  9  retromarginal  denticles. 

Abdomen  2.99  nun  long,  1.98  mm  wide. 
Epigastric  furrow  0.95  mm  from  tracheal 
spiracle,  spiracle  0.90  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.39,  15;  II  1.31,  16;  III  0.75, 
28;  IV  1.44,  14.  Ventral  spination:  tibiae 
I  2-2-2*,  II  1-2-0,  III  1-1-0,  IV  1-1-2; 
metatarsi  as  in  male. 

Epigynum  as  in  Figure  21,  internal  geni- 
talia as  in  Figure  22. 

Natural  history.  Mature  males  have  been 
taken  in  November,  mature  females  from 
early  December  through  late  April.  Speci- 
mens have  been  taken  by  Berlese  funnel 
sampling  of  grape  bark. 

Distribution.  Central  and  southern  Cali- 
fornia ( Map  1 ) . 

Anyphaena  dixiana  (Chamberlin  and 
Woodbury),  new  combination 
Map  1;  Figures  4,  23,  24,  25 

Gaijcnna  dixiana  Chamberlin  and  Woodbun', 
1929,  Proc.  Biol.  Soc.  Washington,  42:  138, 
pi.  1,  fig.  3  (  9  ).  Female  holotype  from  St. 
Ceorge,  Utah,  in  AMNH,  examined.  Roewer, 
1954,  Katalog  der  Araneae,  2:  540  ( G.  dixima 
[sic]).  Bonnet,  1957,  Bibliographia  Araneorum, 
2:  1977. 

Anyphaena  coloradensis  Bryant,  1931,  Psyche, 
38:  112,  pi.  6,  figs.  9,  10,  pi.  7,  figs.  .30,  33 
{  $,  9  ).  Male  holotype,  female  allotNpc  from 
Boulder,  Colorado,  in  MCZ,  examined.   Roewer, 

1954,  Katalog    der   Araneae,    2:    528.     Bonnet, 

1955,  Bibliographia  Araneorum,  2:   343.    NEW 
SYNONYMY. 

Diafinosis.  This  distinctive  species  is 
closest  to  AmjpJuiena  crebrispina,  but  may 
be  quickly  recognized  by  the  spur  borne 
on  the  dorsal  prong  of  the  RTA  of  males 
(Fig.  25)  and  the  greatly  reduced  epigynal 
midpiece  of  females  (Fig.  23). 

Male  (Cochise  Co.,  Arizona).  Coloration 
as  in  Anyphaena  celer  except  that  posterior 
.spiimerets  have  dorsal  surface  sharply  di- 
vided into  dark  brown  lateral  and  pale 
orange  median  halves. 

Total    length    3.85    mm.     Carapace    1.67 


222         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


mm  long,  1.44  mm  wide,  cephalic  width 
0.65  mm,  clypeus  height  0.07  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  6.09, 
PME  0.09,  PLE  0.10;  anterior  eye  row  0.39 
mm  long,  recurved;  posterior  eye  row  0.51 
mm  long,  procurved;  MOQ  length  0.25 
mm,  front  width  0.16  mm,  back  width  0.26 
mm;  eye  interdistances  (mm):  AME- 
AME  0.04,  AME-ALE  0.03,  PME-PME 
0.08,  PME-PLE  0.08,  ALE-PLE  0.05. 

Sternum  0.96  mm  long,  0.76  mm  wide. 
Chelicerae  0.53  mm  long  with  4  promar- 
ginal  teeth  and  6  reti'omarginal  denticles. 
Endites  slightly  invaginated  at  middle. 

Abdomen  2.56  mm  long,  1.49  mm  wide. 
Epigastric  furrow  0.76  mm  from  tracheal 
spiracle,  spiracle  0.76  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  1.69, 
11;  II  1.37,  14;  III  0.81,  28;  IV  1.44,  16. 
Ventral  spination:  tibiae  I,  II  2-2-2*,  III, 
IV  1-2-2;  metatarsi  I,  II  2-2-0,  III  2-0-2, 
IV  2-2-2.  Modifications  of  third  leg  as  in 
A.  celer. 

Palpus  as  in  Figures  4,  25. 

Female  (Cochise  Co.,  Arizona).  Colora- 
tion as  in  male.  Total  length  4.14  mm. 
Carapace  2.03  mm  long,  1.57  mm  wide, 
cephalic  width  0.86  mm,  clypeus  height 
0.09  mm.  Eyes:  diameters  (mm):  AME 
0.05,  ALE  0.08,  PME  0.09,  PLE  0.10;  an- 
terior eye  row  0.43  mm  long,  recurved; 
posterior  eye  row  0.60  mm  long,  procurved; 
MOQ  length  0.26  mm,  front  width  0.20 
mm,  back  width  0.32  mm;  eye  interdis- 
tances (mm):  AME-AME  0.09,  AME- 
ALE  0.05,  PME-PME  0.15,  PME-PLE 
0.09,  ALE-PLE  0.07. 

Sternum  1.15  mm  long,  0.86  mm  wide. 
Chelicerae  0.71  mm  long  with  5  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  2.50  mm  long,  1.69  mm  wide. 
Epigastric  furrow  0.60  mm  from  tracheal 
spiracle,  spiracle  0.67  mm  from  base  of 
.spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.46,  16;  II  1.33,  17;  III  0.94, 
24;  IV  1.49,  17.  Ventral  spination  as  in 
male. 


Epigynum  as  in  Figure  23,  internal  geni- 
talia as  in  Figure  24. 

Natural  history.  Mature  males  have  been 
taken  from  mid- August  through  mid-May, 
mature  females  from  late  September 
through  late  April.  Specimens  have  been 
taken  from  5400  to  9000  feet  (1650-2750 
m),  in  yellow  pine/ oak  and  montane  for- 
ests, in  alfalfa,  under  dead  agave  and  fre- 
quently in  houses. 

Distribution.  Northcentral  Colorado  south 
to  western  Texas,  west  to  southern  Cali- 
fornia (Map  1 ) . 

Anyphaena  judicata  O.  P.-Cambridge 
Map  1;  Figures  5,  26,  33,  34 

Anijphaena  iiidicata  O.  P. -Cambridge,  1896, 
Biologia  Central!  Americana,  Aran.,  1:  203,  pi 
26,  fig.  4  {  S  ).  Male  holotype  from  Omiltemi, 
Guerrero,  Mexico,  in  BMNH,  examined.  F.  O. 
P.-Cambridge,  1900,  Biologia  Centrali  Ameri- 
cana,  Aran.,   2:    96,   pi.   7,   fig.   9,    $.     Roewer, 

1954,  Katalog    der    Araneae,    2:    525.     Bonnet, 

1955,  Bibliographia   Araneormn,   2:    345. 

Diagnosis.  Anyphaena  judicata  is  most 
closely  related  to  an  unnamed  Mexican 
species  (or  group  of  species)  and  has  no 
close  relatives  among  the  species  occur- 
ring north  of  Mexico.  Males  may  be  easily 
recognized  by  the  distinctive  form  of  the 
dorsal  prong  of  the  RTA  (Fig.  26).  The 
female  epigynum  is  closest  to  that  of  A. 
niarginalis,  but  the  midpiece  is  proportion- 
ately shorter  and  wider  and  the  sidepieces 
are  narrower  and  diminish  in  width  an- 
teriorly smoothly,  without  the  sharp  de- 
crease in  width  shown  by  A.  marginalis 
(Fig.  33). 

Male  (Cochise  Co.,  Arizona).  Coloration 
as  in  Anyphaena  celer,  except  that  pos- 
terior spinnerets  have  entire  dorsal  surface 
dark  brown. 

Total  length  3.46  mm.  Carapace  1.76 
mm  long,  1.44  mm  wide,  cephalic  width 
0.68  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.10, 
PME  0.09,  PLE  0.10;  anterior  eye  row  0.40 
mm  long,  recurved;  posterior  eye  row  0.52 
mm  long,  procurved;  MOQ  length  0.26 
mm,    front    width    0.17    mm,    back    width 


Spider  Family  Anyphaenidae  •  Plotnick        223 


0.28  mm;  eye  interdistanees  (mm):  AME- 
AME  0.05,  AME-ALE  0.03,  PME-PME 
0.11,  PME-PLE  0.06,  ALE-PLE  0.04. 

Sternum  0.95  mm  long,  ().6S  mm  wide. 
Chelicerae  0.56  mm  long  with  4  promar- 
ginal  teeth  and  7  retromarginal  denticles. 

Abdomen  1.80  mm  long,  1.15  mm  wide. 
Epigastric  furrow  0.61  mm  from  tracheal 
spiracle,  spiracle  0.63  mm  from  base  of 
.spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.25, 
6;  II  1.93,  8;  III  1.01,  21;  IV  1.66,  11.  Ven- 
tral spination:  tibiae  I  4-2-2*,  II  3-2-2', 
III  1-2-0,  IV  1-1-2;  metatarsi  I,  II,  2-2-0, 
III  2-0-2;  IV  1-2-2.  Femur  III  unmodi- 
fied. Tibia  III  ventral  spine  1  on  retrolat- 
eral  side  missing,  ventral  spine  2  thickened, 
cone-like.  Coxae  I,  II  and  III  ( but  not  IV ) 
with  a  small  number  of  short,  thick  setae. 
Coxae  III  with  a  tiibercule. 

Palpus  as  in  Figures  5,  26. 

Female  (Cochise  Co.,  Arizona).  Colora- 
tion as  in  male. 

Total  length  4.72  mm.  Carapace  1.76 
mm  long,  1.37  mm  wide,  cephalic  width 
0.81  mm,  clypeus  height  0.06  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.10, 
PME  0.10,  PLE  0.10;  anterior  eye  row  0.44 
mm  long,  recurved;  posterior  eye  row  0.60 
mm  long,  procurved;  MOQ  length  0.29 
mm,  front  width  0.21  mm,  back  width  0.32 
mm;  eye  interdistanees  (mm):  AME- 
AME  0.07,  AME-ALE  0.03,  PME-PME 
0.13,  PME-PLE  0.09,  ALE-PLE  0.05. 

Sternum  0.97  mm  long,  0.77  mm  wide. 
Chelicerae  0.58  mm  long  with  teeth  as  in 
male. 

Abdomen  3.13  mm  long,  2.09  mm  wide. 
Epigastric  furrow  1.21  mm  from  tracheal 
spiracle,  spiracle  1.31  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.69,  11;  II  1.31,  14;  III  0.88, 
23;  IV  1.66,  12.  Ventral  spination  as  in 
male. 

Epigynum  as  in  Figure  33,  internal  geni- 
talia as  in  Figure  34. 

Natural  history.  Mature  males  have  been 
taken  from  mid-June  through  mid-August, 


mature  females  from  late  March  to  Novem- 
ber, most  ill  July  and  August.  Specimens 
have  been  taken  from  5100  to  8000  feet 
(1550-2450  m),  by  sweeping  and  under 
rocks. 

Distribution.  Arizona  south  to  Guerrero, 
Mexico  (Map  1). 

Anyptiaena  marginalis  (Banks), 
new  combination 
Map  1;  Figures  6,  27,  36,  43 

Gayeima  marginalis  Banks,  1901,  Proc.  Acad. 
Natur.  Sci.  Philadelphia,  53:  574,  pi.  2.3,  fig. 
22  (  9  ).  Female  holotype  from  Beulali,  San 
Miguel  Co.,  New  Mexico,  was  probabl>'  de- 
posited in  the  MCZ  along  with  the  other  types 
from  this  paper  but  was  not  found  by  Bryant 
when  the  MCZ  t>pes  were  cataloged;  lost, 
presumed  destroyed.  Roewer,  1954,  Katalog 
der  Araneae,  2:  540.  Bonnet,  19.57,  Biblio- 
graphia  Araneorum,  2:  1978. 

Diagnosis.  Amjphaena  marginalis  is  most 
closely  related  to  A.  hespar,  both  species 
having  a  simple  embolus  and  elongated 
conductor.  Males  of  A.  marginalis  (Fig. 
27),  however,  do  not  have  the  spine-like 
dorsal  prong  of  the  RTA  of  A.  hespar,  and 
females  of  A.  marginalis  (Fig.  36)  do  not 
have  the  conspicuous  bulge  in  the  epigynal 
midpiece  which  characterizes  A.  hespar 
females. 

Male  (Graham  Co.,  Arizona).  Colora- 
tion as  in  Anyphaena  celer. 

Total  length  3.78  mm.  Carapace  1.98 
mm  long,  1.60  mm  wide,  cephalic  width 
0.72  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.10, 
PME  0.08,  PLE  0.10;  anterior  eye  row  0.40 
mm  long,  straight;  posterior  eye  row  0.55 
mm  long,  prociuved;  xVlOQ  length  0.20 
mm,  front  width  0.17  mm,  back  width  0.28 
mm;  eye  interdistanees  (mm):  AME- 
AME  0.05,  AME-ALE  0.02,  PME-PME 
0.11,  PME-PLE  0.08,  ALE-PLE  0.04. 

Sternum  1.13  mm  long,  0.81  mm  wide. 
Chelicerae  0.54  mm  long  with  5  promar- 
ginal  teeth  and  6  retromarginal  denticles. 

Abdomen  2.00  mm  long,  1.33  mm  wide. 
Epigastric  furrow  0.52  mm   from   traclieal 


224 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


spiracle,    spiracle   0.59   mm    from   base   of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  1.67, 
13;  II  1.35,  16;  III  1.03,  26;  IV  1.62,  14. 
Ventral  spination:  tibiae  I  4-2-2*,  II  2-2- 
2*,  III,  IV  1-2-2;  metatarsi  I,  II  2-2-0,  III 
2-0-2,  IV  2-2-2.  Femur  III  unmodified. 
Tibia  III  ventral  spine  1  on  retrolateral 
side  missing.    Coxae  unmodified. 

Palpus  as  in  Figures  6,  27. 

Female  (Graham  Co.,  Arizona).  Colora- 
tion as  in  male  of  A.  celer. 

Total  length  4.26  mm.  Carapace  2.11 
mm  long,  1.55  mm  wide,  cephalic  width 
0.86  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.10, 
PME  0.11,  PLE  0.10;  anterior  eye  row  0.44 
mm  long,  straight;  posterior  eye  row  0.64 
mm  long,  procurved;  MOQ  length  0.30 
mm,  front  width  0.19  mm,  back  width  0.33 
mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.03,  PME-PME 
0.12,  PME-PLE  0.09,  ALE-PLE  0.06. 

Sternum  1.05  mm  long,  0.80  mm  wide. 
Chelicerae  0.65  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  2.52  mm  long,  1.53  mm  wide. 
Epigastric  furrow  0.67  mm  from  tracheal 
spiracle,  spiracle  0.68  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.44,  18;  II  1.21,  21;  III  0.99, 
25;  IV  1.60,  17.  Ventral  spination  as  in 
male  except  tibia  III  1-1-2. 

Epigynum  as  in  Figure  36,  internal  geni- 
talia as  in  Figure  43. 

'Natural  history.  Mature  males  have  been 
taken  from  late  August  through  late  May, 
mature  females  in  all  months  except  Janu- 
ary and  October.  Specimens  have  been 
taken  from  6000  to  9300  feet  (1850-2850 
m),  in  yellow  pine/ oak  forests  and  under 
rocks.  I  found  this  species  in  great  abun- 
dance by  sorting  pine  litter  at  Rustler's 
Park  in  the  Chiricahua  Mountains  of  south- 
eastern Arizona  in  August  1972. 

Distribution.  Arizona,  New  Mexico  and 
Colorado  (Map  1). 


Anyphaena  hespar  new  species 
Map  1;  Figures  16,  32,  42,  49 

Types.  Male  holotype,  female  paratype 
from  Bear  Canyon,  Santa  Catalina  Moun- 
tains, Pima  Co.,  Arizona,  8  December  1968 
(Karl  Stephan),  deposited  in  AMNH.  Male 
and  female  paratypes  from  Pima  Co.,  Ari- 
zona, deposited  in  MCZ.  The  specific 
name  is  an  arbitrary  combination  of  letters. 

Diagnosis.  Anyphaena  hespar  is  most 
closely  related  to  A.  marginalis.  Males  of 
the  former  may  be  distinguished  by  the 
spine-like  dorsal  prong  of  their  RTA  (Fig. 
32),  females  by  the  conspicuous  bulge  in 
their  epigynal  midpiece  (Fig.  42). 

Male  (Pima  Co.,  Arizona).  Coloration 
as  in  Anyphaena  celer. 

Total  length  3.13  mm.  Carapace  1.62 
mm  long,  1.31  mm  wide,  cephalic  width 
0.59  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.08, 
PxME  0.08,  PLE  0.08;  anterior  eye  row 
0.33  mm  long,  straight;  posterior  eye  row 
0.45  mm  long,  procurved;  MOQ  length  0.19 
mm,  front  width  0.14  mm,  back  width  0.24 
mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.03,  PME-PME 
0.08,  PME-PLE  0.07,  ALE-PLE  0.04. 

Sternum  0.95  mm  long,  0.79  mm  wide. 
Chelicerae  0.39  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  1.80  mm  long,  1.10  mm  wide. 
Epigastric  furrow  0.56  mm  from  tracheal 
spiracle,  spiracle  0.56  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  1.31, 
17;  II  1.08,  21;  III  0.81,  28;  IV  1.39,  18. 
Ventral  spination:  tibiae  I  4-2-2*,  II  3- 
2-2*,  III  1-2-2,  IV  2-2-2;  metatarsi  I,  II 
2-2-0,  III  2-0-2,  IV  2-2-2.  Femur  III  un- 
modified. Tibia  III  ventral  spine  1  on  retro- 
lateral  side  missing,  spine  2  thickened, 
cone-like.   Coxae  unmodified. 

Palpus  as  in  Figures  16,  32. 

Female  (Pima  Co.,  Arizona).  Colora- 
tion as  in  male  of  A.  celer. 

Total  length  3.06  mm.  Carapace  1.55 
mm  long,  1.26  mm  wide,  cephalic  width 
0.67  mm,  clypeus  height  0.06  mm.    Eyes: 


Spider  Family  Anvphaenidak  •  Plafuick        225 


diameters  (mm):  AME  0.05,  ALE  O.OS, 
PME  0.08,  PLE  O.OS;  anterior  eye  row  0.33 
mm  long,  .straight;  posterior  eye  row  0.49 
mm  long,  procurved;  MOQ  length  0.20 
mm,  front  width  0.14  mm,  back  width  0.26 
mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.03,  PME-PME 
0.10,  PME-PLE  0.07,  ALE-PLE  0.04. 

Sternum  1.04  mm  long,  0.70  mm  wide. 
Chelicerae  0.47  mm  long  with  teeth  as  in 
male. 

Abdomen  1.85  mm  long,  1.08  mm  wide. 
Epigastric  furrow  0.49  mm  from  tracheal 
spiracle;  spiracle  0.41  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.13,  20;  II  0.92,  25;  III  0.72, 
33;  IV  1.26,  18.  Ventral  spination:  tibiae 
I,  II  4-2-2^  III  1-1-2,  IV  1-2-2;  meta- 
tarsi as  in  male. 

Epigynum  as  in  Figure  42,  Internal  geni- 
talia as  in  Figure  49. 

Natural  history.  Mature  males  and  fe- 
males have  been  taken  from  late  October 
through  early  April.  Specimens  have  been 
taken  from  leaf  litter  and  under  rocks. 

DistriJmtion.  Southeastern  Arizona  (Map 

1)- 

Anyphaena  rita  new  species 
Map  1;  Figures  7,  28,  37,  44 

Types.  Male  holotype,  female  paratype 
from  Bear  Canyon,  Santa  Catalina  Moun- 
tains, Pima  Co.,  Arizona,  8  December  1968 
(Karl  Stephan),  deposited  in  AMNH.  Male 
and  female  paratypes  from  Pima  Co.,  Ari- 
zona, deposited  in  MCZ.  The  .specific 
name  is  a  noun  in  apposition  derived  from 
the  Santa  Rita  Mountains,  where  the 
species  is  abundant. 

Diagnosis.  Anypliaemi  rita  is  most  closely 
related  to  A.  cochise,  both  species  having 
a  conspicuously  enlarged  region  of  the 
embolus  and  a  slightly  recurved  tip  of  the 
median  apophysis.  Males  of  A.  rita  (Fig. 
28)  may  be  distingui.shed  by  their  smaller 
size  and  by  the  differences  in  the  dorsal 
prong  of  the  RTA.  Females  of  A.  cochise 
are  unknown,  l)ut  the  epigynum  of  A.  rita. 


with  its  extremely  broad  midpiece,  is  quite 
distinctive  (Fig.  37). 

Male  (Pima  Co.,  Arizona).  Colorati(;n 
as  in  Anyphaena  celer. 

Total  length  4.10  mm.  Carapace  1.94 
mm  long,  1.60  mm  wide,  cephalic  width 
0.67  mm,  clypeus  height  0.07  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.08, 
PME  0.09,  PLE  0.09;  anterior  eye  row  0.36 
mm  long,  recurved;  posterior  eye  row  0.53 
mm  long,  procurved;  MOQ  length  0.22 
mm,  front  width  0.15  mm,  back  width  0.27 
mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.03,  PME-PME 
0.09,  PME-PLE  0.09,  ALE-PLE  0.05. 

Sternum  1.13  mm  long,  0.77  mm  wide. 
Chelicerae  0.50  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  2.30  mm  long,  1.26  mm  wide. 
Epigastric  furrow  0.67  mm  from  tracheal 
spiracle,  spiracle  0.65  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  1.55, 
14;  II  1.39,  16;  III  0.97,  23;  IV  1.62,  14. 
Ventral  spination:  tibiae  I  4-2-2*,  II 
3-2-2*,  III,  IV  2-2-2;  metatarsi  I,  II 
2-2-0,  III  2-0-2,  IV  2-2-2.  Femur  III  un- 
modified. Tibia  III  ventral  spines  not 
thickened.  Coxae  III  and  IV  with  only  a 
few  short  thick  setae. 

Palpus  as  in  Figures  7,  28. 

Female  (Pima  Co.,  Arizona).  Coloration 
as  in  male  of  A.  celer. 

Total  length  5.04  mm.  Carapace  2.05 
mm  long,  1.53  mm  wide,  cephalic  width 
1.03  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  6.10, 
PME  0.10,  PLE  0.11;  anterior  eye  row  0.41 
mm  long,  recurved;  posterior  eye  row  0.58 
mm  long,  procurved;  MOQ  length  0.32 
mm,  front  width  0.18  mm,  back  width 
0.29  mm;  eye  interdistances  (mm):  AME- 
AME  0.07,'  AME-ALE  0.03,  PME-PME 
0.09,  PME-PLE  0.10,  ALE-PLE  0.08. 

Sternum  1.13  mm  long,  0.81  mm  wide. 
Chelicerae  0.67  mm  long  with  teeth  as  in 
male. 

Abdomen  2.75  mm  long,  1.94  mm  wide. 
Epigastric  furrow   1.06   mm   from   tracheal 


226         Bulletin  Museum  of  Comparative  Zoologij,  Vol.   146,  No.   4 


spiracle,  spiracle  0.95  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.48,  15;  II  1.26,  18;  III 
1.03,  21;  IV  1.58,  17.  Ventral  spination  as 
in  male  except  tibiae  III,  IV  1-2-2,  meta- 
tarsi IV  2-1-2. 

Epigynum  as  in  Figure  37,  internal 
genitalia  as  in  Figure  44. 

Natural  history.  Mature  males  have  been 
taken  from  mid-October  through  late 
March,  mature  females  from  early  June 
tlirough  early  February.  Specimens  have 
been  taken  from  4000  to  6800  feet.  ( 1200- 
2075  m),  in  oak/ grassland  and  under  rocks. 

Distribution.  Arizona  to  Chihuahua, 
Mexico  (Map  1). 

Anyphaena  cochise  new  species 
Map  1;  Figures  13,  35 

Types.  Male  holotype  from  Rustlers 
Park,  8600  ft.  (2625  m),  Chiricahua  Moun- 
tains, Cochise  Co.,  Arizona,  9  September 
1950  (W.  J.  Gertsch),  deposited  in  AMNH. 
Male  paratype  from  Cochise  Co.,  Arizona, 
deposited  in  MCZ.  The  specific  name  is  a 
noun  in  apposition  and  refers  to  the  type 
locality. 

Diapiosis.  Anypliaena  cochise  is  most 
closely  related  to  A.  vita,  but  the  dorsal 
prong  of  the  RTA  is  relatively  longer  in 
A.  cochise  (Fig.  35).  Females  of  this 
species  are  unknown. 

Male  (Cochise  Co.,  Arizona).  Colora- 
tion as  in  Anyphaena  celer. 

Total  length  5.44  mm.  Carapace  2.52 
mm  long,  2.09  mm  wide,  cephalic  width 
0.88  mm,  clypeus  height  0.14  mm.  Eyes: 
diameters  (mm):  AME  0.09,  ALE  6.13, 
PME  0.13,  PLE  0.13;  anterior  eye  row 
0.53  mm  long,  straight;  posterior  eye  row 
0.75  mm  long,  procurved;  MOQ  length 
0.30  mm,  front  width  0.23  mm,  back  width 
0.40  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.04,  PME-PME 
0.14,  PME-PLE  0.11,  ALE-PLE  0.06. 

Sternum  1.44  mm  long,   1.08  mm  wide. 


Chelicerae  0.75  mm  long  with  4  promar- 
ginal  teeth  and  7  retromarginal  denticles. 

Abdomen  3.38  mm  long,  1.94  mm  wide. 
Epigastric  fvuTow  0.92  mm  from  tracheal 
spiracle,  spiracle  1.03  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.32, 
12;  II  2.05,  13;  III  1.39,  20;  IV  2.14,  14. 
Ventral  spination:  tibiae  I  4-2-2*,  II  2-2- 
2*,  III  1-2-2,  IV  2-2-2;  metatarsi  I,  II  2- 
2-0,  III  2-0-2,  IV  2-2-2.  Femur  III  un- 
modified. Tibia  III  ventral  spine  1  on  ret- 
rolateral  side  thickened  slightly.  All  coxae 
with  a  few  scattered  short  thick  setae. 

Palpus  as  in  Figures  13,  35. 

Female.  Unknown. 

Natural  history.  Mature  males  have  been 
taken  in  early  September  at  8600  feet 
(2625  m). 

Distribution.  Known  only  from  the  type 
locality  (Map  1). 

Anyphaena  autumna  new  species 
IVIap  1;  Figures  8,  29,  39,  45 

Types.  Male  holotype,  female  paratype 
from  Rustler  Camp,  Chiricahua  Mountains, 
Cochise  Co.,  Arizona,  9  September  1950 
(W.  J.  Gertsch),  deposited  in  AMNH. 
Male  and  female  paratypes  from  Cochise 
and  Graham  Co.,  Arizona,  deposited  in 
MCZ.  The  specific  name  refers  to  the 
season  of  collection. 

Diagno.sis.  Anyphaena  autumna  is  un- 
likely to  be  confused  with  any  other  spe- 
cies. The  long  recurved  hook  on  the  RTA 
and  the  peculiar  form  of  the  tip  of  the 
median  apophysis  are  mil  ike  any  other 
species  (Figs.  8,  29).  The  epigynum  is 
closest  to  that  of  A.  gibboides,  but  the  mid- 
piece  has  a  characteristic  constriction  near 
its  midpoint  ( Fig.  39 ) . 

Male  (Cochise  Co.,  Arizona).  Colora- 
tion as  in  Anyphaena  celer,  though  the 
paramedian  bands  on  the  carapace  are 
darker  and  wider  than  in  that  species. 

Total  length  5.51  mm.  Carapace  2.50 
mm  long,  1.98  mm  wide,  cephalic  width 
1.03  mm,  clypeus  height  0.12  mm.    Eyes: 


Spider  Family  Anyphaenidae  •  Platnick        227 


diameters  (nini):  AME  0.09,  ALE  0.12, 
PME  0.12,  PLE  0.13;  anterior  eye  row 
0.55  mm  long,  recnrved;  posterior  eye  row 
0.75  mm  long,  proeun'ed;  MOQ  length 
0.30  mm,  front  width  0.26  mm,  back  width 
0.38  mm;  eye  interdistances  (mm):  AME- 
AME  0.08,  AME-ALE  0.05,  PME-PME 
0.15,  PME-PLE  0.11,  ALE-PLE  0.06. 

Sternnm  1.46  mm  long,  1.08  mm  wide. 
Chelicerae  0.79  mm  long  with  4  promar- 
ginal  teeth  and  9  retromarginal  denticles. 

Abdomen  3.20  mm  long,  2.16  mm  wide. 
Epigastric  furrow  1.04  mm  from  tracheal 
spiracle,  spiracle  1.06  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.16, 
13;  II  1.93,  15;  III  1.39,  22;  IV  2.16,  14. 
Ventral  spination:  tibiae  I  2-2-2,  II,  III, 
IV  1-2-2;  metatarsi  I,  II  2-2-0,  III  2-0-2, 
I\^  2-2-2.    Third  legs  unmodified. 

Palpus  as  in  Figures  8,  29. 

Female  (Cochise  Co.,  Arizona).  Colora- 
tion as  in  male. 

Total  length  6.41  mm.  Carapace  2.34 
mm  long,  1.87  mm  wide,  cephalic  width 
1.12  mm,  clypeus  height  0.12  mm.  Eyes: 
diameters  (mm):  AME  0.10,  ALE  0.13, 
PME  0.13,  PLE  0.13;  anterior  eye  row 
0.59  mm  long,  recurved;  posterior  eye  row 
0.70  mm  long,  procurved;  MOQ  length 
0.33  mm,  front  width  0.27  mm,  back  width 
0.42  mm;  eye  interdistances  (mm):  AME- 
AME  0.06,'  AME-ALE  0.03,  PME-PME 
0.17,  PME-PLE  0.12,  ALE-PLE  0.07. 

Sternum  1.42  mm  long,  1.08  mm  wide. 
Chelicerae  0.99  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  3.96  mm  long,  2.63  mm  wide. 
Epigastric  furrow  1.33  mm  from  tracheal 
spiracle,  spiracle  1.33  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.75,  16;  II  1.60,  18;  III  1.10, 
2.5;  IV  1.89,  15.  Ventral  spination:  tibiae 
I  4-4-2,  II  2-4-2,  III  1-1-2,  IV  1-2-2; 
metatarsi  as  in  male. 

Epigynum  as  in  Figure  39,  internal  geni- 
talia as  in  Figure  45. 

Natural  history.    Mature   males   and   fe- 


males have  been  taken  in  August  and  Sep- 
tember. Specimens  have  be(Mi  taken  at 
8200  fec>t  (2500  m).  I  collected  a  few  im- 
matiue  males  (which  matured  in  the  labo- 
ratory) of  this  .species  in  pine  litter  in  the 
Chiricahua  Mountains,  Arizona,  where  ma- 
ture A.  mar^inalis  were  extremely  abun- 
dant. 

Dustribution.  Southeastern  Arizona  (Map 

1)- 

Anyphaena  gibboides  new  species 
Map  1;  Figures  14,  30,  40,  46 

Types.  Male  holotype,  female  paratype 
from  City  Creek  Canyon,  Salt  Lake  Co., 
Utah,  22  May  1943  (Wilton  Ivie),  depos- 
ited in  AMNH.  Male  and  female  para- 
t)^es  from  Lake  Co.,  Oregon,  deposited  in 
MCZ.  The  specific  name  is  an  arbitrary 
combination  of  letters. 

Diagnosis.  Anyphaena  gihJ)oi(Ies  is  a 
distinctive  species.  Males  have  a  sharply 
pointed  median  apophysis  and  serrate 
RTA  which  will  separate  them  from  the 
other  known  species  (Figs.  14,  30).  The 
epigynum  is  closest  to  that  of  A.  autumna, 
but  lacks  the  constriction  of  the  midpiece 
found  in  that  species  ( Fig.  40 ) . 

Male  (Salt  Lake  Co.,  Utah).  Coloration 
as  in  Anyphaena  celer. 

Total  length  3.31  mm.  Carapace  1.60 
mm  long,  1.28  mm  wide,  cephalic  width 
0.54  mm,  clypeus  height  0.07  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.08, 
PME  0.08,  PLE  0.08;  anterior  eye  row  0.34 
mm  long,  straight;  posterior  eye  row  0.48 
mm  long,  procurved;  MOQ  length  0.23 
mm,  front  width  0.15  mm,  back  width  0.24 
mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.08,  PME-PME 
0.08,  PME-PLE  0.07,  ALE-PLE  0.05. 

Sternum  0.85  mm  long,  0.72  mm  wide. 
Chelicerae  0.49  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  1.94  mm  long,  1.24  mm  wide. 
Epigastric  furrow  0.58  mm  from  tracheal 
spiracle,  spiracle  0.59  mm  from  base  of 
spinnerets. 


228         BuUetm  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


Tibial  lengths  (mm)  and  indices:  I  1.33, 
17;  II  1.24,  19;  III  0.99,  23;  IV  1.47,  16. 
Ventral  spination:  tibiae  I  2-2-0,  II  1-2-0, 
III  2-2-0,  IV  2-2-2;  metatarsi  I,  II  2-2-0, 
III  2-1-2,  IV  2-2-2.  Modifications  of  third 
leg  as  in  A.  celer  save  that  all  coxae  have 
clumps  of  short  thick  setae. 

Palpus  as  in  Figures  14,  30. 

Female  (Salt  Lake  Co.,  Utah).  Colora- 
tion as  in  male  of  A.  celer. 

Total  length  3.74  mm.  Carapace  1.75 
mm  long,  1.35  mm  wide,  cephalic  width 
0.83  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.09, 
PME  O.OS,  PLE  0.08;  anterior  eye  row 
0.41  mm  long,  recurved;  posterior  eye  row 
0.57  mm  long,  procurved;  MOQ  length 
0.24  mm,  front  width  0.18  mm,  back  width 
0.28  mm;  eye  interdistances  (mm):  AME- 
AME  0.06,  AME-ALE  0.03,  PME-PME 
0.12,  PME-PLE  0.09,  ALE-PLE  0.06. 

Sternum  1.19  mm  long,  0.83  mm  wide. 
Chelicerae  0.62  mm  long  with  4  promar- 
ginal  teeth  and  6  retromarginal  denticles. 

Abdomen  2.36  mm  long,  1..39  mm  wide. 
Epigastric  furrow  0.72  mm  from  tracheal 
spiracle,  spiracle  0.70  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.39,  18;  II  1.24,  20;  III 
0.75,  27;  IV  1.39,  18.  Ventral  spination: 
tibiae  I  2-2-0,  II,  III  1-2-0,  IV  1-2-2; 
metatarsi  I,  II  2-2-0,  III  2-0-2,  IV  2-2-2. 

Epigynum  as  in  Figure  40,  internal  geni- 
talia as  in  Figure  46. 

Natural  history.  Mature  males  and  fe- 
males have  been  taken  in  late  May  and 
June.  Habitat  data  is  lacking. 

Distribution.  Northern  Utah  west  to 
southeastern  Oregon  ( Map  1 ) . 

Anyphaena  catalina  new  species 
Map  1;  Figures  15,  31,  41,  48 

Types.  Male  holotype,  female  paratype 
from  Mt.  Lemon,  Santa  Catalina  Moun- 
tains, Pima  Co.,  Arizona,  13  July  1916  (  F. 
E.  Lutz),  deposited  in  AMNH.  Male  and 
female  paratypes  from  Pima  Co.,  Arizona, 


and  Mexico,  Mexico,  deposited  in  MCZ. 
The  specific  name  is  a  noun  in  apposition 
and  refers  to  the  type  locality. 

Diagnosis.  Anyphaena  catalina  is  most 
closely  related  to  A.  arbida,  though  males 
of  A.  catalina  may  be  readily  distinguished 
by  their  recurved  retrolateral  tegular 
apophyses  (Figs.  15,  31).  Females  of  A. 
arbida  are  unknown;  those  of  A.  catalina 
may  be  recognized  by  the  epigynal  hood 
being  roughly  equal  in  size  to  the  epigynal 
midpiece  ( Fig.  41 ) . 

Male  (Pima  Co.,  Arizona).  Coloration 
as  in  Anyphena  celer. 

Total"  length  3.53  mm.  Carapace  1.78 
mm  long,  1.42  mm  wide,  cephalic  width 
0.72  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.09, 
PME  0.08,  PLE  0.09;  anterior  eye  row 
0.40  mm  long,  recurved;  posterior  eye  row 
0.51  mm  long,  procurved;  MOQ  length 
0.21  mm,  front  width  0.17  mm,  back  width 
0.26  mm;  eye  interdistances  (mm):  AME- 
AME  0.07,  AME-ALE  0.04,  PME-PME 
0.09,  PME-PLE  0.08,  ALE-PLE  0.04. 

Sternum  0.90  mm  long,  0.70  mm  wide. 
Chelicerae  0.56  mm  long  with  4  promar- 
ginal  teeth  and  6  retromarginal  denticles. 

Abdomen  1.85  mm  long,  0.90  mm  wide. 
Epigastric  furrow  0.61  mm  from  tracheal 
spiracle,  spiracle  0.65  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.07, 
8;  II  1.94,  9;  III  1.08,  23;  IV  1.80,  10.  Ven- 
tral spination:  tibiae  I  4-2-2*,  II  2-2-2*, 
III,  IV  2-2-2;  metatarsi  I,  II  2-2-0,  III  2- 
0-2,  IV  2-2-2.  Modifications  of  third  leg 
as  in  A.  celer  save  that  femur  III  lacks  short 
thick  setae  and  all  coxae  bear  clumps  of 
them. 

Palpus  as  in  Figures  15,  31. 

Female  (Pima  Co.,  Arizona).  Coloration 
as  in  male  of  A.  celer. 

Total  length  4.57  mm.  Carapace  1.84 
mm  long,  1.42  mm  wide,  cephalic  width 
0.94  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.09, 
PME  0.09,  PLE  0.09;  anterior  eye  row 
0.47  mm  long,  recui-ved;  posterior  eye  row 


Spideu  pAisriLY  Anyphaemdak  •  Pintnick        229 


0.63  mm  long,  prociirved;  MOQ  length  0.26 
mm,  front  width  0.22  mm,  back  widtli  0.33 
mm;  eve  interdi.stances  (mm):  AME- 
AME  6.0S,  AME-ALE  0.04,  PME-PME 
0.15,  PME-PLE  0.11,  ALE-PLE  0.07. 

Sternum  1.01  mm  long,  0.85  mm  wide. 
Chelicerae  0.68  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticle.s. 

Abdomen  2.74  nnn  long,  1.85  mm  wide. 
Epigastric  furrow  0.86  mm  from  tracheal 
.spiracle,  spiracle  0.94  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.51,  14;  II  1.33,  17;  III  0.94, 
23;  IV  1.48,  16.  Ventral  spination:  tibiae  I, 
II  2-2-2*,  III  1-2-2,  IV  2-2-2;  metatarsi 
as  in  male. 

Epigynum  as  in  Figure  41,  internal  geni- 
talia as  in  Figure  48. 

Natural  Justory.  Mature  males  and  fe- 
males have  been  taken  in  July  and  August. 
Specimens  have  been  taken  at  7500  feet 
(2300  m)  in  yellow  pine/ oak  and  douglas 
fir/ white  fir  forests. 

Distribution.  Southeastern  Arizona  south 
to  central  Mexico  (Map  1). 

Anyphaena  arbida  new  species 
Map  1;  Figures  17,  38 

Types.  Male  holotype  from  Carr  Can- 
yon, Huachuca  Mountains,  Cochise  Co., 
Arizona,  26  August  1950  (M.  A.  Cazier), 
deposited  in  AMNH.  Male  paratype  from 
Cochise  Co.,  Arizona,  deposited  in  MCZ. 
The  specific  name  is  an  arbitrary  combina- 
tion of  letters. 

Diagnosis.  AnypJiaena  arhida  is  most 
closely  related  to  A.  catalina.  Males  of  the 
former  (Figs.  17,  38)  lack  the  recurved 
retrolateral  tegular  apophysis  of  A.  cata- 
lina; females  of  A.  arbida  are  unknown. 

Male  (Cochise  Co.,  Arizona).  Colora- 
tion as  in  AnypJiaena  celer,  except  that 
posterior  spinnerets  are  as  in  A.  dixiana. 

Total  length  6.95  mm.  Carapace  3.28 
mm  long,  2.41  mm  wide,  cephalic  width 
1.22  mm,  clypeus  height  0.14  mm.  Eyes: 
diameter    (mm):    AME    0.11,    ALE    0.13, 


PME  0.13,  PLE  0.15;  anterior  eye  row 
0.64  mm  long,  recurved;  posterior  eye  row 
0.89  mm  long,  procurved;  MOQ  lengtii 
0.43  mm,  front  width  0.31  mm,  back  width 
0.44  mm;  eye  interdistances  (mm):  AME- 
AME  0.09,  AME-ALE  0.04,  PME-PME 
0.18,  PME-PLE  0.14,  ALE-PLE  0.09. 

Sternum  1.62  mm  long,  1.33  mm  wide. 
Chelicerae  1.30  mm  long  with  4  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  3.71  mm  long,  2.16  mm  wide. 
Epigastric  furrow  1.08  mm  from  tracheal 
spiracle,  .spiracle  1.12  mm  from  base  of 
spinnerets.  Spinnerets  surrounded  by  a 
clump  of  unusually  long  setae. 

Tibial  lengths  (mm)  and  indices:  I  6.88, 
5;  II  3.35,  10;  III  2.20,  16;  IV  3.35,  10. 
Ventral  spination:  tibiae  I  4-2-2*,  II  3- 
2-2*,  III,  IV  2-2-0;  metatarsi  I,  II  2-2-0, 
III,  IV  2-2-2.   Third  legs  unmodified. 

Palpus  as  in  Figures  17,  38. 

Female.  Unknown. 

Natural  history.  Mature  males  have  been 
collected  in  August.    Habitat  data  is  lack- 


ing- 


1). 


Di.stribution.  Cochise  Co.,  Arizona  (Map 


Anyphaena  pectorosa  Group 


Diagnosis.  The  pectorosa  group  is  closely 
related  to  the  pacifica  group,  but  males 
may  be  distinguished  by  the  spins  on  their 
coxae  (Figs.  59-62).  Females  have  the 
epigynum  on  a  characteristic  sclerotized 
plate  (Figs.  74,  77,  79)  and  simple  .sper- 
mathecae  (Figs.  75,  78,  80). 

Description.  Total  length  4.5-6.5  mm. 
Carapace  longer  than  wide,  narrowed  in 
front  to  less  than  half  its  maximum  width 
in  males,  to  slightly  more  than  half  its 
maximum  width  in  females.  Clypeus  height 
more  than  1.5  times  the  diameter  of  an  an- 
terior median  eye.  Posterior  median,  pos- 
terior lateral  and  anterior  lateral  eyes  sub- 
equal  in  size,  almost  twice  the  diameter  of 
anterior  medians.  Procurved  posterior  eye 
row  longer  than  slightly  recur\xKl  anterior 
row.      Median    ocular    (juadrangle    almost 


230         Bulletin  Museum  of  Comparative  Zoolofi.ij,  Vol.   146,  No.   4 


twice  as  wide  in  back  as  in  front.  Anterior 
median  eyes  separated  l:)y  sliglitly  less  than 
their  diameter,  sHghtly  closer  to  anterior 
laterals  than  to  each  other.  Posterior  me- 
dians separated  by  slightly  more  than  their 
diameter,  slightly  closer  to  posterior  lat- 
erals. Anterior  laterals  separated  by  their 
radins  from  posterior  laterals.  Sternum 
longer  than  wide,  with  a  low  hirsute  knob 
behind  its  middle  in  some  males.  Chelic- 
erae  with  4  promarginal  teeth  and  7-9 
retromarginal  denticles.  Abdomen  longer 
than  wide,  tiacheal  spiracle  midway  be- 
tween epigastric  furrow  and  base  of  spin- 
nerets. Leg  formula  1423.  Metatarsi  I 
and  11  with  two  pairs  of  ventral  spines. 
Males  with  coxae  II  bearing  round  knobs, 
coxae  III  and  IV  bearing  spurs.  Palpus 
with  an  elongated  median  apophysis,  en- 
larged conductor  and  inconspicuous  embo- 
lus. Retrolateral  tibial  apophysis  short. 
Epigynum  on  a  sclerotized  plate,  without 
a  hood.    Two  simple  spermathecae. 

Variation.  The  species  in  this  group 
show  little  intraspecific  variation,  individ- 
ual or  geographical,  in  size,  structure  or 
coloration. 

Key  to  Species 

la.  Coxae  III  of  males  with  posterior  spur  bifid 
(Fiffs.  59,  61,  62);  sternum  of  males  with  a 
low  hirsute  knob  behind  middle;  sclerotized 
epigynal  plate  wider  posteriorly  than  an- 
teriorly   (Figs.   74,   79)    2 

lb.  Coxae  III  of  males  with  posterior  spur  not 
bifid  ( Fig.  60 ) ;  sternum  of  males  without  a 
low  hirsute  knob  behind  middle;  sclerotized 
epigynal  plate  wider  anteriorly  than  pos- 
teriorly   (Fig.   77)    fratema 

2a.  Distal  tip  of  palpal  median  apophysis  bent 
sharply  towards  cymbium  (Figs.  55,  58); 
sclerotized  epigynal  plate  with  pronounced 
posterolateral    corners    ( Fig.    74 )    3 

2b.  Distal  tip  of  palpal  median  apophysis  not 
bent  sharply  towards  cymbium  ( Fig. 
57 ) ;  sclerotized  epigynal  plate  without  pro- 


nounced posterolateral  corners  (Fig.  79) 
alaclma 

3a.  Distal  tip  of  palpal  median  apophysis  meet- 
ing the  recessed,  dorsal  branch  of  tlie  apoph- 
ysis ( Fig.  55 ) ;  sclerotized  epigynal  plate 
with  pronounced  posterolateral  comers 
( Fig.  74 )    pectorosa 

3b.  Distal  tip  of  palpal  median  apophysis  not 
meeting  the  recessed,  dorsal  branch  of  the 
apophysis  (Fig.  58);  females  unknown  __7flc?:a 

Anyphaena  pectorosa  L.  Koch 
Map  2;  Figures  51,  55,  59,  74,  75 

Anyphaena  pectorosa  L.  Koch,  1866,  Arachn.  Fam. 
Drass.,  198,  pi.  8,  figs.  131,  132  {  $).  Male 
holotype  from  Baltimore,  Maryland,  in  BMNH, 
examined.  Bryant,  1931,  Psyche,  38:  110,  pi.  6, 
fig.  5,  $ .  Chickering,  1939,  Pap.  Michigan 
Acad.  Sci.,  24:  51,  figs.  5-8,  $,9.  Comstock, 
1940,  Spider  Book,  rev.  ed.,  p.  577,  fig.  636,  9 . 
Kaston,  1948,  Bull.  Connecticut  Geol.  Natur. 
Hist.  Surv.,  70:  408,  figs.  1453,  1477-1480, 
$,  9.  Roewer,  1954,  Katalog  der  Araneae,  2: 
529.  Bonnet,  1955,  Bibliographia  Araneorum, 
2:  346. 

Aniiphaena  calcarata  Emerton,  1890,  Trans.  Con- 
necticut Acad.  Sci.,  8:  187,  pi.  6,  figs.  3-3d  (  $, 
9  ).  Male  holotype,  female  allotype  from  West 
Haven,  Connecticut,  in  MCZ,  examined.  Emer- 
ton, 1902,  Common  Spiders,  p.  12,  figs.  42,  43, 
$,  9. 

Gaijenna  calcarata,  Banks,  1910,  Bull.  U.S.  Nat. 
Mus.,  72:  13. 

Gaijenna  pectorosa,  Comstock,  1912,  Spider  Book, 
p.  563  (in  part),  fig.  636,   9    (not  fig.  637). 

Diagnosis.  Anyphaena  pectorosa  is  closest 
to  A.  alachua,  but  may  readily  be  distin- 
guished from  it  by  the  highly  curved  me- 
dian apophysis  of  males  (Fig.  55)  and  the 
pronounced  posterolateral  corners  of  the 
sclerotized  epigvnal  plate  of  females  (Fig. 
74). 

Male  (Fairfax  Co.,  Virginia).  Total 
length  5.40  mm.  Carapace  2.43  mm  long, 
1.98  mm  wide,  cephalic  width  0.88  mm, 
clypeus  height  0.11  mm,  yellow  with  thin 
dark  border  and  two  dark  paramedian 
longitudinal  bands.   Eyes:  diameters  (mm): 


Plate  5 
Figures  51--54.    Left  palpi,  ventral  view.    Figures  55-58.    Left  palpi,  retrolateral  view.    Figures  59-62. 
ventral  view.     51,  55,  59.    Anyptiaena  pectorosa  L.  Koch.     52,  56,  60.    Anyphaena  fraterna  (Banks). 
Anyphaena  alachua  new  species.     54,  58,  62.    Anyphaena  lacka  new  species. 


Male  coxae, 
53,  57,  61. 


Spider  I'^aafily  Anyphaenidak  •  Plaliuck        231 


232         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  4 


AME  0.06,  ALE  0.11,  PME  0.11,  PLE  0.12; 
anterior  e\e  row  0.48  mm  long,  slightly  re- 
cur^•ed;  posterior  eye  row  0.65  mm  long, 
procurved;  MOQ  length  0.28  mm,  front 
width  0.20  mm,  back  width  0.35  mm;  eye 
interdistances  (mm):  AME-AME  0.07, 
AME-ALE  0.04,  PME-PME  0.14,  PME- 
PLE  0.13,  ALE-PLE  0.05. 

Sternum  1.35  mm  long,  1.01  mm  wide, 
pale  yellow  with  translucent  border,  dark- 
ened extensions  to  coxae  and  a  low  hirsute 
knob  behind  middle.  Chelicerae  0.73  mm 
long  with  4  promarginal  teeth  and  7  retro- 
marginal  denticles,  pale  yellow  with  boss 
outlined  in  gray.  Labium  and  endites  yel- 
low, darkest  proximally.  Endites  slightly 
invaginated  at  middle. 

Abdomen  3.15  mm  long,  1.67  mm  wide, 
pale  white  with  transverse  rows  of  dark 
markings,  venter  pale.  Epigastric  furrow 
1.01  mm  from  tracheal  spiracle,  spiracle 
1.06  mm  from  base  of  spinnerets. 

Legs  pale  yellow  with  distal  segments 
darkest.  Tibial  lengths  (mm)  and  indices: 
I  3.10,  7;  II  2.52,  9;  III  1.82,  16;  IV  2.56,  10. 
Ventral  spination:  tibiae  I  2-2-1,  II-IV  2- 
2-2;  metatarsi  I,  II  2-2-0,  III  2-0-2,  IV  2- 
2-2.  Coxae  II,  III  and  IV  modified  as  in 
Figure  59. 

Palpus  as  in  Figures  51,  55. 

Female  (Fairfax  Co.,  Virginia).  Colora- 
tion as  in  male. 

Total  length  5.44  mm.  Carapace  2.41 
mm  long,  1.91  mm  wide,  cephalic  width 
0.97  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.12, 
PME  0.11,  PLE  0.12;  anterior  eye  row 
0.52  mm  long,  recurved;  posterior  eye  row 
0.71  mm  long,  procurved;  MOQ  length 
0.33  mm,  front  width  0.20  mm,  back  width 
0.37  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.04,  PME-PME 
0.15,  PME-PLE  0.10,  ALE-PLE  0.07. 

Sternum  1.31  mm  long,  1.06  mm  wide, 
without  hirsute  knob.  Chelicerae  0.72  mm 
long  with  4  promarginal  teeth  and  8  retro- 
marginal  denticles. 

Abdomen  3.10  mm  long,  1.76  mm  wide. 
Epigastric  furrow  0.70  mm  from  tracheal 


spiracle,  spiracle  1.22  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  2.41,  11;  II  2.05,  13;  III  1.44, 
19;  IV  2.20,  12.  Ventral  spination:  tibiae 
I,  II  2-2-0,  III,  IV  1-2-1;  metatarsi  I,  II 
2-2-0,  III,  IV  2-2-2. 

Epigynum  as  in  Figure  74,  internal  geni- 
talia as  in  Figure  75. 

Natural  history.  Mature  males  have  been 
taken  from  mid-April  through  early  Sep- 
tember, mature  females  from  mid-April 
through  mid-August.  Specimens  have  been 
taken  by  sweeping  foliage,  in  Malaise  and 
pitfall  ti^ips,  and  under  rocks.  Egg  cases 
taken  with  females  contained  65-95  eggs. 

Distribution.  New  England  west  to 
Michigan,  south  to  western  Florida  and 
eastern  Texas  ( Map  2 ) . 

Anyphaena  atachua  new  species 
Map  2;  Figures  53,  57,  61,  79,  80 

Types.  Male  holotype,  female  paratype 
from  west  of  Gainesville,  Alachua  Co., 
Florida,  18  April  1938  (Willis  J.  Certsch), 
deposited  in  AMNH.  Male  and  female 
paratypes  from  Alachua  Co.,  Florida,  de- 
posited in  MCZ.  The  specific  name  is  a 
noun  in  apposition  and  refers  to  the  type 
locality. 

Diagnosis.  Anyphaena  alachua  is  closest 
to  A.  pectorosa  but  the  median  apophysis 
is  not  highly  curved  (Fig.  57)  and  the 
epigynal  plate  lacks  pronounced  postero- 
lateral corners  (Fig.  79). 

Male  (Alachua  Co.,  Florida).  Colora- 
tion as  in  Anyphaena  pectorosa. 

Total  length  4.90  mm.  Carapace  2.41 
mm  long,  2.01  mm  wide,  cephalic  width 
0.79  mm,  clypeus  height  0.13  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.12, 
PME  0.12,  PLE  0.13;  anterior  eye  row 
0.51  mm  long,  slightly  recurved;  posterior 
eye  row  0.70  mm  long,  procurved;  MOQ 
length  0.30  mm,  front  width  0.22  mm,  back 
width  0.36  mm;  eye  interdistances  (mm): 
AME-AME  0.07,  AME-ALE  0.04,  PME- 
PME  0.12,  PME-PLE  0.11,  ALE-PLE  0.06. 


Spider  Family  Anvi'haemdae  •  Plalnick        233 


Sternum  1.26  mm  long,  1.01  mm  wide, 
with  low  hirsute  knob  Ix^hind  middle. 
Chelicerae  0.76  mm  long  with  4  promar- 
ginal  teeth  r.nd  9  retromarginal  denticle.s. 

Abdomen  2.48  mm  long,  1.48  mm  wide. 
Epigastric  furrow  0.76  mm  from  tracheal 
spiracle,  spiracle  0.8.3  mm  from  base  of 
spinnerc>ts. 

Tibial  lengtlis  (nun)  and  indices:  I  2.77, 
10;  II  2.27,^11;  III  1.44,  22;  IV  1.94.  14. 
\Vntral  spination:  tibiae  I,  II  2-2-0,  III 
1-2-2,  IV  2-2-2;  metatarsi  I,  II  2-2-0,  III 
2-0-2,  IV  2-2-2.  Coxae  II,  III  and  I\' 
modified  as  in  Figure  61. 
Palpus  as  in  Figures  53,  57. 
Female  (Alachua  Co.,  Florida).  Colora- 
tion as  in  male  of  A.  pectorosa. 

Total  length  6.17  mm.  Carapace  2.45 
mm  long,  1.80  mm  wide,  cephalic  width 
0.94  mm,  clypeus  height  0.12  mm.  Eyes: 
diameters  (mm):  AME  0.08,  ALE  0.13, 
PME  0.12,  PLE  0.13;  anterior  eye  row 
0.57  mm  long,  slightly  recurved;  posterior 
eye  row  0.73  mm  long,  procui-ved;  MOQ 
IcMigth  0.30  mm,  front  width  0.22  mm,  back 
width  0.40  mm;  eye  interdistances  (mm): 
AME-AME  0.07,  AME-ALE  0.04,  PME- 
PME  0.15,  PME-PLE  0.11,  ALE-PLE 
0.07. 

Sternum  1.35  mm  long,  1.08  mm  wide, 
without  hirsute  knob.  Clielicerae  0.84  mm 
long  with  teeth  as  in  male. 

Abdomen  3.53  mm  long,  2.02  mm  wide. 
Epigastric  furrow  1.10  mm  from  tracheal 
spiracle,  spiracle  1.21  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  2.30,  13;  II  1.91,  14;  III  1.31, 
22;  IV  2.09,  13.  Ventral  spination  as  in 
male  save  metatarsi  III  2-2-2. 

Epigynum  as  in  Figure  79,  internal  geni- 
talia as  in  Figure  80. 

Natural  history.  Mature  males  have  been 
taken  in  late  April  and  early  May,  mature 
females  from  late  March  through  mid-May, 
by  sweeping. 

Distribution.  Known  only  from  Florida 
(Map  2). 


Anyphaena  lacka  new  species 
Map  2;  Figures  54,  58,  62 

Type.  Male  liolotxpe  from  Lake  Corpus 
Christi  State  Park,  southwest  of  Mathis, 
San  Patricio  Co.,  Texas,  28  Jvme  1962  (J.  A. 
Beatty),  deposited  in  MCZ.  The  specific 
name  is  an  arbitrary  combination  of  letters. 
Dia<i,nosis.  Anyphaena  lacka  is  most 
closely  reflated  to  A.  alachua  ])ut  has  a  dis- 
tinct point  on  the  tip  of  the  median  apoph- 
ysis (Fig.  58).  Females  of  A.  lacka  are 
unknown. 

Male  (San  Patricio  Co.,  Texas).   Colora- 
tion as  in  Anyphaena  pectorosa. 

Total  length  4.61  mm.  Carapace  2.05 
mm  long,  1.69  mm  wide,  cephalic  width 
0.79  mm,  clypeus  height  0.12  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.12, 
PME  0.11,  PLE  0.11;  anterior  eye  row 
0.47  mm  long,  slightly  recurved;  posterior 
eye  row  0.61  mm  long,  procurved;  MOQ 
length  0.26  mm,  front  width  0.19  mm,  back 
width  0.32  mm;  eye  interdistances  (mm): 
AME-AME  0.05,  AME-ALE  0.03,  PME- 
PME  0.11,  PME-PLE  0.08,  ALE-PLE  0.04. 
Sternum  1.24  mm  long,  0.90  mm  wide, 
with  low  hirsute  knob  behind  middle. 
Chelicerae  0.64  mm  long  with  4  promar- 
ginal  teeth  and  9  retromarginal  denticles. 

Abdomen  2.41  mm  long,  1.33  mm  wide. 
Epigastric  furrow  0.74  mm  from  tracheal 
spiracle,  spiracle  0.90  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.38, 
9;  II  1.91,  12;  III  1.32,  30;  IV  1.93,  12. 
Ventral  spination:  tibiae  I,  II  2-2-0,  III, 
IV  1-2-2;  metatarsi  I,  II  2-2-0,  III  2-0-2, 
IV  2-2-2.  Coxae  II,  III,  and  W  modified 
as  in  Figure  62. 

Palpus  as  in  Figures  54,  58. 
Female.  LTnknown. 

Natural  history  and  distribution.  Known 
only  from  the  type  specimen. 

Anyphaena  fraterna  (Banks) 

iVlap  2;  Figures  52,  56,  60,  77,  78 

.\]iil})hacna     coiispcrsa     KeyserlinK.     1887,     Verh. 
zool.   bot.   Cos.   Wien,   37:    453,   pi.   6,   fig.   23 


234         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


(  $  ).  Female  holotype  from  Bee  Spring,  Ken- 
tucky, in  MCZ,  examined;  preoccupied  by  Any- 
phaena  conspersa  Simon,   1878. 

Gaijenna  fratema  Banks,  1896,  Trans.  Amer.  Ent. 
Soc,  23:  63.  Male  holotype  from  Sea  Cliff, 
New  York,  in  MCZ,  examined. 

Amjphaena  fratema,  Simon,  1897,  Hist.  Natur. 
Araign.,  2:  96.  Bryant,  1931,  Psyche,  38:  110, 
pi.  6,  fig.  6,  pi.  8,  fig.  23,  $,  9.  Comstock, 
1940,  Spider  Book,  rev.  ed.,  p.  577,  fig.  637,  $  . 
Kaston,  1948,  Bull.  Connecticut  Geol.  Natur. 
Hist.  Surv.,  70:  408,  figs.  1454-1456,  $,  9. 
Roewer,  1954,  Katalog  der  Araneae,  2:  529. 
Bonnet,  1955,  Bibliographia  Araneonnn,  2:  344. 

Sillus  consperstis,  Petrunkevitch,  1911,  Bull.  Amer. 
Mus.  Natur.  Hist.,  29:  511. 

Gayenna  pectorosa,  Comstock,  1912,  Spider  Book, 
p.  563  (in  part),  fig.  637,  $. 

Diagnosis.  Anyphaena  fratema  is  a  dis- 
tinctive species  easily  recognized  by  the 
long  and  narrow  median  apophysis  of 
males  (Fig.  52)  and  by  the  female's  epigy- 
nal  plate  being  wider  anteriorly  than  pos- 
teriorly (Fig.  77). 

Male  (Hall  Co.,  Georgia).  Coloration 
as  in  Anyphaena  pectorosa. 

Total  length  4.93  mm.  Carapace  2.23 
mm  long,  1.85  mm  wide,  cephalic  width 
0.81  mm,  clypeus  height  0.10  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.12, 
PME  0.11,  PLE  0.11;  anterior  eye  row  0.45 
mm  long,  slightly  recurved;  posterior  eye 
row  0.64  mm  long,  procurved;  MOQ  length 
0.30  mm,  front  width  0.18  mm,  back  width 
0.33  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.04,  FME-PiME 
0.12,  PME-PLE  0.08,  ALE-PLE  0.05. 

Sternum  1.28  mm  long,  0.99  mm  wide, 
without  hirsute  knob.  Chelicerae  0.59  mm 
long  with  4  promarginal  teeth  and  9  retro- 
marginal  denticles. 

Abdomen  2.83  mm  long,  1.60  mm  wide. 
Epigastric  furrow  0.97  mm  from  tracheal 
spiracle,  spiracle  0.85  mm  from  base  of 
spinnerets. 

Legs  with  scattered  dark  spots.  Tibial 
lengths  (mm)  and  indices:  I  2.60,  8;  II 
2.16,  11;  III  1.52,  16;  IV  2.47,  10.  Ventral 
.spination:  tibiae  I,  II  2-2-0,  III  1-2-2,  IV 
2-2-2;  metatarsi  I,  II  2-2-0,  III,  IV  2-2-2. 
Coxae  II,  III  and  IV  modified  as  in  Figure 
60. 


Palpus  as  in  Figures  52,  56. 

Female  (Hall  Co.,  Georgia).  Coloration 
as  in  male  of  A.  pectorosa. 

Total  length  5.00  mm.  Carapace  2.32 
mm  long,  1.80  mm  wide,  cephalic  width 
0.94  mm,  clypeus  height  0.10  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.11, 
PME  0.11,  PLE  0.11;  anterior  eye  row 
0.49  mm  long,  recurved;  posterior  eye  row 
0.69  mm  long,  procui-ved;  MOQ  length 
0.27  mm,  front  width  0.18  mm,  back  width 
0.36  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,'  AME-ALE  0.04,  PME-PME 
0.15,  PME-PLE  0.09,  ALE-PLE  0.05. 

Sternum  1.28  mm  long,  1.04  mm  wide. 
Chelicerae  0.75  mm  long  with  teeth  as  in 
male. 

Abdomen  2.97  mm  long,  1.71  mm  wide. 
Epigastric  furrow  0.85  mm  from  tracheal 
spiracle,  spiracle  0.85  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  2.29,  11;  II  1.89,  13;  III  1.30, 
19;  IV  2.16,  13.  Ventral  spination  as  in 
male. 

Epigynum  as  in  Figure  77,  internal  geni- 
talia as  in  Figure  78. 

Natural  history.  Mature  males  have  been 
taken  from  late  March  through  early  July, 
mature  females  from  late  March  through 
late  August.  Specimens  have  been  taken 
by  sweeping  foliage,  in  Malaise  and  pitfall 
traps,  and  under  logs.  I  collected  this  spe- 
cies in  great  abundance  by  sweeping 
honeysuckle  (Lonicera  sp.)  at  night  in 
southern  West  Virginia  during  June  1971. 

Distribution.  Southern  New  York  west 
to  eastern  Kansas,  south  to  western  Florida 
and  eastern  Texas  (Map  2). 

Anyphaena  pacifica  Group 

Diagnosis.  The  pacifica  group  is  closely 
related  to  the  pectorosa  group  and  appears 
to  displace  it  in  the  western  United  States. 
The  males  have  similarly  short  retrolateral 
tibial  apophyses  (Figs.  69-71),  but  paci- 
fica group  males  lack  the  coxal  spurs  char- 
acteristic  of  the   pectorosa   group,   though 


Spider  Family  ANYPiiAiixiDAt;  •  Flatnick        235 


<: 


Anyphoena  pectorosa    o\  -*  > 


I 


------v*- 


i£ 


Anyphoena  fraterna    .«\  -^> 


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Anyphaenc 

pacificQ 

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Anyphoena 

colifornicQ    - 

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Anyphoena 

locko 

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Anyphoena  aperta    ' 

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Map  2.     Distributions  of  Anyphaena  alachua,  A.  aperta,  A.  californica,  A.  fraterna,  A.  gertschi,  A.  lacka,  A.  pacif- 
ica  and  A.  pectorosa. 


males  of  Anyphaena  gertschi  have  rounded 
knobs  on  the  coxae.  Females  lack  the 
sclerotized  epigynal  plates  found  in  the 
pectorosa  group,  but  have  a  lightly  sclero- 
tized atrivun-like  area  posteromedially 
(Figs.  66,  67,  72)  and  long,  sometimes 
coiling,  ducts  (Figs.  68,  73,  76). 

Description.  Total  length  4-6  mm.  Cara- 
pace longer  than  wide,  narrowed  in  front 
by  at  least  one-third  of  its  maximum  width, 
often  by  more  than  half.  Clypeus  height 
roughly  equal  to  anterior  median  eye  diam- 
eter. All  eyes  subequal  in  size.  Procurved 
posterior  eye  row  longer  than  slightly  re- 
curved anterior  eye  row.  Median  ocular 
quadrangle  longer  than  wide  in  front, 
wider  in  back  than  long.  Anterior  median 
eyes  separated  by  less  than  their  diameter, 
much  closer  to  anterior  laterals  than  to 
each  other.  Posterior  medians  separated 
by  more  than  their  diameter,  much  closer 
to  posterior  laterals.  Anterior  laterals  sepa- 
rated by  slightly  more  than  their  radius 
from    posterior    laterals.     Sternum    longer 


than  wide,  without  a  hirsute  knob.  Chelic- 
erae  with  3  promarginal  teeth  and  6-9 
retromarginal  denticles.  Abdomen  longer 
than  wide,  tracheal  spiracle  midway  be- 
tween epigastric  furrow  and  base  of  spin- 
nerets. Leg  formula  1423.  Metatarsi  I  and 
II  with  two  pairs  of  ventral  spines.  Males 
with  legs  unmodified.  (A.  pacifica  and  A. 
californica)  or  with  coxae  bearing  round 
knobs  and  femora  II  and  III  bearing 
patches  of  short  stiff  setae  ventrally  (A. 
gertschi).  Palpus  with  an  elongated  me- 
dian apophysis,  enlarged  conductor  and 
inconspicuous  embolus.  Retrolateral  tibial 
apophysis  short.  Epigyiuim  not  on  a  scle- 
rotized plate,  without  a  hood,  with  a  more 
or  less  pronoimced  atrium-like  lightly 
sclerotized  area  posteromedially.  Internal 
genitalia  with  long  ducts  that  coil  in  some 
species. 

Variation.  Two  species  in  this  group,  A. 
pacifica  and  A.  californica,  show  a  great 
deal  of  xariation  in  genitalic  structure.  In 
both  species  the  shape  oi  the  tip  of  the  pal- 


236         Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.   4 


pal  median  apophysis  and  the  coihng  of 
the  epigynal  ducts  are  strikingly  variable, 
and  it  was  initially  thought  that  many 
species  were  involved.  Three  sources  of 
evidence,  however,  have  indicated  other- 
wise. First,  many  females  are  found  in 
which  the  ducts  on  one  side  of  the  epigy- 
num  coil  differently  from  those  on  the 
other  side.  Secondly,  when  many  speci- 
mens are  taken  together  at  one  locality  on 
a  single  da}",  several  variants  are  often 
found.  Finally,  the  retrolateral  tibial 
apophysis,  which  usually  provides  excellent 
diagnostic  characters  in  anyphaenids,  is 
stable  within  the  species  as  they  are  de- 
fined here.  Until  such  time  as  biological 
evidence  on  the  breeding  habits  of  these 
spiders  can  be  obtained,  it  seems  best  to 
consider  both  A.  pacifica  and  A.  califorjuco 
as  widespread,  variable  species. 

Key  to  Species 

la.  Retrolateral  tibial  apophysis  (RTA)  without 
a  dorsal  process  (Fig.  69).  Median  apoph- 
ysis with  a  deep  invagination  below  tip 
giving  the  tip  a  chelate  appearance  (Fig. 
65).  Epigyninii  with  large  wing-shaped 
paramedian  flaps   ( Fig.  72 )   gertschi 

lb.  Retrolateral  tibial  apophysis  (RTA)  with  a 
dorsal  process  (Figs.  70,  71).  Median 
apophysis  without  a  deep  invagination  be- 
low tip  (Figs.  63,  64).  Epigynum  with- 
out large  wing-shaped  paramedian  flaps 
(Figs.   66,   67)    2 

2a.  Dorsal  process  of  RTA  short,  located  dis- 
tally  (Fig.  70).  Median  apophysis  narrow- 
ing gradually  towards  tip  (Fig.  63).  In- 
ternal ducts  with  many  coils  (Fig.  68)  — _ 
—     pacifica 

2b.  Dorsal  process  of  RTA  long,  located  prox- 
inially  (Fig.  71).  Median  apophysis  nar- 
rowing abruptly  towards  tip  (Fig.  64).  In- 
ternal ducts  without  many  coils  (Fig.  73) 
— californica 


Anyphaena  pacifica  (Banks) 
Map  2;  Figures  63,  66,  68,  70 

Gaijcnna  pacifica  Banks,  1896,  Trans.  Amer.  Ent. 
Soc,  23:  63.  Female  holot>'pe  from  Olympia, 
Washington,  in  MCZ,  examined. 

Anyphaena  pacifica,  Simon,  1897,  Hist.  Natur. 
Araign.,  2:  96.  Bryant,  1931,  Psyche,  38:  115, 
pi.  8,  fig.  36,  ?.  Levi  and  Levi,  1951,  Zoo- 
logica  (New  York),  36:  228,  Tig.  25,  $.  Roe- 
wer,  1954,  Katalog  der  Araneae,  2:  529.  Bon- 
net, 1955,  Bibliographia  Araneorum,  2:  346. 

Anyphaena  mundella  Chamberlin,  1920,  Pomona 
Coll.  J.  Ent.  Zool.,  12:  12,  pi.  5,  fig.  3  (  9 ,  not 
$,  =  Aysha  incursa) .  Female  holotype  from 
Claremont,  California,  in  MCZ,  examined. 
Bryant,  1931,  Psyche,  38:  120  (sub  Aysha  de- 
cepta  [sic] ).  Roewer,  1954,  Katalog  der  Araneae 
2:  534  (sub  Aysha  decepta  [sic]).  Bonnet, 
1955,  Bibliographia  Araneorum,  2:  836  (sub 
Aysha  decepta   [sic]).    NEW  SYNONYMY. 

Anyphaena  intermontana  Chamberlin,  1920, 
Canad.  Ent.,  52:  200,  fig.  22-6  (  $  ).  Female 
holotype  from  Mill  Creek,  Salt  Lake  Co.,  Utali, 
in  MCZ,  examined.  Bryant,  1931,  Psyche,  38: 
114  (sub  Anyphaena  californica  [sic]).  Roe- 
wer, 1954,  Katalog  der  Araneae,  2:  528  (sub 
Anyphaena  californica  [sic]).  Bonnet,  1955, 
Bibliographia  Araneorum,  2:  343  (sub  Any- 
phaena californica    [sic]).    NEW  SYNONYMY. 

Gayenna  saniuana  Chamberlin  and  Gertsch,  1928, 
Proc.  Biol.  Soc.  Wash.,  41:  185.  Male  holotype 
from  Verdure,  San  Juan  Co.,  Utah,  in  AMNH, 
examined.  Roewer,  1954,  Katalog  der  Araneae, 
2:  540.   NEW  SYNONYMY. 

Anyphaena  saniuana,  Bryant,  1931,  Psyche,  38: 
107.  Bonnet,  1955,  Bibliographia  Araneorum, 
2:  347. 

Anyphaena  pomona  Chamberlin  and  Ivie,  1941, 
Bull.  Univ.  Utah,  Biol.,  6:  23,  pi.  2,  fig.  16 
(  9  ).  Female  holotype  from  Mill  Creek,  Te- 
hama Co.,  California,  in  AMNH,  examined. 
Roewer,  1954,  Katalog  der  Araneae,  2:  529. 
NEW  SYNONYMY. 

Gayenna  jollensis  Schenkel,  1950,  Verb.  Naturf. 
Ges.  Basel,  61:  77,  fig.  27  (  9  ).  Female  holo- 
type from  La  Jolla,  California,  in  Naturhistor- 
isches  Museum,  Basel,  examined.  Roewer,  1954, 
Katalog  der  Araneae,  2:  540.  NEW  SYN- 
ONYMY. 


Plate  6 
Figures  63-65.  Left  palpi,  ventral  view.  Figures  69-71.  Left  palpal  tibiae,  retrolateral  view.  Figures  66,  67,  72, 
74,  77,  79.  Epigyna,  ventral  view.  Figures  68,  73,  75,  76,  78,  80.  Internal  genitalia,  dorsal  view.  63,  66,  68,  70. 
Anyptiaena  pacifica  {Banks).  64,67,71,73.  Anyptiaena  californica  {Banks).  65,69,72,76.  Anyphaena  gertschi 
new  species.  74,  75.  Anyphaena  pectorosa  L.  Koch.  77,  78.  Anyphaena  fraterna  (Banks).  79,  80.  Anyphaena 
alachua  new  species. 


Spider  Family  Anyphaenidae  •  Plalnick        237 


68 


^V 


?) 


78 


•^ 


£S^' 


238         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


Diagnosis.  Amjphaena  pacifica  is  closest 
to  A.  californica,  but  males  may  be  distin- 
guished by  the  short,  distal,  dorsal  process 
of  the  retrolateral  tibial  apophysis  (Fig. 
70)  and  the  gradually  narrowing  tip  of  the 
median  apophysis  (Fig.  63),  while  females 
have  distinctive  highly  coiled  internal 
ducts  (Fig.  68).  Variation  in  this  species 
is  discussed  above. 

Male  (El  Dorado  Co.,  California).  Total 
length  5.18  mm.  Carapace  2.34  mm  long, 
1.94  mm  wide,  cephalic  width  0.86  mm, 
clypeus  height  0.12  mm,  pale  orange  with 
thin  dark  border  and  two  dark  paramedian 
longitudinal  bands.  Eyes:  diameters  (mm): 
AME  0.09,  ALE  0.12,  PME  0.10,  PLE  0.11; 
anterior  eye  row  0.51  mm  long,  slightly 
procurved;  posterior  eye  row  0.69  mm  long, 
procurved;  MOQ  length  0.28  mm,  front 
width  0.24  mm,  back  width  0.34  mm;  eye 
interdistances  (mm):  AME-AME  0.07, 
AME-ALE  0.03,  PME-PME  0.14,  PME- 
PLE  0.10,  ALE-PLE  0.05. 

Sternum  1.49  mm  long,  1.04  mm  wide, 
pale  orange  with  darker  border.  Chelicerae 
0.67  mm  long  with  3  promarginal  teeth 
and  8  retromarginal  denticles,  dark  orange- 
brown  proximally,  pale  orange  distally, 
with  boss  outlined  in  gray.  Labium  and 
endites  orange,  darkest  proximally.  En- 
dites  slightly  invaginated  at  middle. 

Abdomen  2.81  mm  long,  1.69  mm  wide, 
reddish-brown  throughout.  Epigastric  fur- 
row 0.85  mm  from  tracheal  spiracle,  spira- 
cle 0.92  mm  from  base  of  spinnerets. 

Legs  pale  orange,  unmodified.  Tibial 
lengths  (mm)  and  indices:  I  2.11,  12;  II 
1.87,  13;  III  1.44,  20;  IV  2.07,  15.  Ventral 
spination:  tibiae  I,  II  2-2-0,  III  1-2-2,  IV 
2-2-2;  metatarsi  I,  II  2-2-0,  III,  IV  2-2-2. 

Palpus  as  in  Figures  63,  70. 

Female  (Mono  Co.,  California).  Color- 
ation as  in  male. 

Total  length  5.39  mm.  Carapace  2.34 
mm  long,  1.62  mm  wide,  cephalic  width 
0.94  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.10,  ALE  0.12, 
PME  0.11,  PLE  0.11;  anterior  eye  row  0.51 
mm  long,  slightly  recurved;  posterior  eye 


row  0.73  mm  long,  procurved;  MOQ  length 
0.29  mm,  front  width  0.25  mm,  back  width 
0.36  mm;  eye  interdistances  (mm):  AME- 
AME  0.06,  AME-ALE  0.03,  PME-PME 
0.15,  PME-PLE  0.10,  ALE-PLE  0.07. 

Sternum  1.44  mm  long,  1.01  mm  wide. 
Chelicerae  0.71  mm  long  with  teeth  as  in 
male. 

Abdomen  3.02  mm  long,  1.69  mm  wide. 
Epigastric  furrow  0.81  mm  from  tracheal 
spiracle,  spiracle  0.86  mm  from  base  of 
spinerets. 

Tibial  lengths  (mm)  and  indices:  I  1.84, 
15;  II  1.71,  15;  III  1.39,  19;  IV  2.07,  13. 
Ventral  spination  as  in  male  save  tibiae 
III  1-1-2  and  IV  1-2-2. 

Epigynum  as  in  Figure  66,  internal  geni- 
talia as  in  Figure  68. 

Natural  history.  Mature  males  have  been 
taken  from  late  February  through  late 
July,  mature  females  year  round.  Speci- 
mens have  been  taken  in  montane  forests, 
in  pitfall  traps,  under  rocks  and  commonly 
in  houses. 

Distribution.  Western  North  America 
from  British  Columbia  south  to  California, 
Ai-izona  and  New  Mexico  (Map  2). 

Anyphaena  californica  (Banks) 
Map  2;  Figures  64,  67,  71,  73 

Gaijenna  californica  Banks,  1904,  Proc.  California 
Acad.  Sci.,  3:  338,  pi.  38,  fig.  2(9).  Female 
holotype  from  Palo  Alto,  California,  in  MCZ, 
examined. 

Amiphaena  mens  Chamberlin,  1920,  Pomona  Coll. 
J.  Ent.  Zoo!.,  12:  11,  pi.  5,  fig.  \  {$).  Male 
holotype  from  Claremont,  California,  in  MCZ, 
examined.  Bryant,  1931,  Psyche,  38:  113. 
Roewer,  1954,  Katalog  der  Araneae,  2:  529. 
Bonnet,  1955,  Bibliographia  Araneorum,  2:  347. 
NEW  SYNONYMY. 

Anyphaena  californica,  Bryant,  1931,  Psyche,  38: 
114.  Roewer,  1954,  Katalog  der  Araneae,  2: 
528.  Bonnet,  1955,  Bibliographia  Araneorum, 
2:  343. 

Diagnosis.  Amjphaena  californica  is  most 
closely  related  to  A.  pacifica,  but  males 
have  a  long,  proximal,  dorsal  process  on 
the  retrolateral  tibial  apophysis  (Fig.  71) 
and  an  abruptly  narrowed  tip  of  the  me- 


Spider  Family  Anyphaenidae  •  Phifnick 


239 


diaii  apopliysis  (Fig.  64),  while  the  inter- 
nal dncts  of  the  female  are  not  highly 
coiled  (Fig.  73).  Variation  in  this  species 
is  discnssed  above. 

Male  (San  Diego  Co.,  California).  Col- 
oration as  in  AnypJiaena  pacifica  except 
that  the  abdomen  is  pale  white  with  trans- 
verse rows  of  dark  markings. 

Total  length  4.68  mm.  Carapace  2.21 
mm  long,  1.78  mm  wide,  cephalic  width 
0.68  mm,  clypeus  height  0.07  mm.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  6.09, 
PME  0.10,  PLE  0.11;  anterior  eye  row 
0.43  mm  long,  recm-ved;  posterior  eye  row 
0.59  mm  long,  procurved;  MOQ  length 
0.30  mm,  front  width  0.20  mm,  back  width 
0.32  mm;  eye  interdistances  (mm):  AME- 
AME  0.06,  AME-ALE  0.04,  PME-PME 
0.13,  PME-PLE  0.11,  ALE-PLE  0.07. 

Sternum  1.31  mm  long,  0.90  mm  wide. 
Chelicerae  0.60  mm  long  with  3  promar- 
ginal  teeth  and  8  retromarginal  denticles. 

Abdomen  2.97  mm  long,  1.34  mm  wide. 
Epigastric  furrow  0.79  mm  from  tracheal 
spiracle,  spiracle  0.85  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  3.28, 
5;  II  3.20,  7;  III  2.27,  8;  IV  2.93,  7.  Ventral 
spination:  tibiae  I  2-2-0,  II  2-2-2,  III  1- 
1-2,  IV  1-2-2;  metatarsi  I,  II  2-2-0,  III, 
I\'  2-2-2. 

Palpus  as  in  Figures  64,  71. 

Female  (Humboldt  Co.,  CaHfornia). 
Coloration  as  in  male. 

Total  length  5.98  mm.  Carapace  2.56 
mm  long,  1.91  mm  wide,  cephalic  width 
1.03  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.09,  ALE  0.11, 
PME  0.12,  PLE  0.12;  anterior  eye  row  0.52 
mm  long,  recurved;  posterior  eye  row  0.69 
mm  long,  procurved;  MOQ  length  0.35 
mm,  front  width  0.25  mm,  back  width  0.37 
mm;  eye  interdistances  (mm):  AME- 
AME  0.07,  AME-ALE  0.04,  PME-PME 
0.13,  PME-PLE  0.10,  ALE-PLE  0.06. 

Sternum  1.44  mm  long,  1.08  mm  wide. 
Chelicerae  0.86  mm  long  with  3  promar- 
ginal  teeth  and  9  retromarginal  denticles. 

Abdomen  3.64  mm  long,  2.43  mm  wide. 


Epigastric  furrow  1.15  mm  from  tracheal 
spiracle,  spiracle  1.33  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.16, 
12;  II  1.87,  13;  III  1.30,  19;  IV  2.06,  14. 
Ventral  spination  as  in  male  except  tibiae 
II  1-2-0. 

Epigynum  as  in  Figure  67,  internal  geni- 
talia as  in  Figure  73. 

Natural  Jii.story.  Mature  males  have  been 
taken  from  early  March  through  mid-July, 
mature  females  from  mid-March  through 
mid-November.  Specimens  have  Ix^en 
taken  in  redwood  forests,  on  citrus  trees 
and  in  houses. 

Distribution.  Oregon  and  California 
(Map  2). 

Anyphaena  gertschi  new  species 
Map  2;  Figures  65,  69,  72,  76 

Types.  Male  holotype,  female  paratype 
from  Bluff,  San  Juan  Co.,  Utah,  11  May 
1933  (Wilton  Ivie),  deposited  in  AMNH. 
Male  and  female  paratypes  from  Emery 
Co.,  Utah,  deposited  in  MCZ.  The  specific 
name  is  a  patronym  in  honor  of  Willis  J. 
Gertsch,  who  first  recognized  the  species 
as  new. 

Diagnosis.  Anyphaena  gertschi  is  a  dis- 
tinctive species  easily  recognized  by  the 
chelate  appearance  of  the  tip  of  the  me- 
dian apophysis  of  males  (Fig.  65)  and 
the  large  wing-shaped  paramedian  flaps 
on  the  female  epigynum  (Fig.  72). 

Male  (Emery  Co.,  Utah).  Coloration  as 
in  Anyphaena  pacifica  except  that  cara- 
pace has  paramedian  bands  only  vaguely 
indicated  and  abdomen  is  pale  yellow 
throughout. 

Total  length  4.00  mm.  Carapace  1.85 
mm  long,  1.42  mm  wide,  cephalic  width 
0.92  mm,  clypeus  height  0.14  mm.  Eyes: 
diameters  (mm):  AME  0.09,  ALE  6.09, 
PME  0.09,  PLE  0.09;  anterior  eye  row 
0.45  mm  long,  slightly  recurv'ed;  pos- 
terior eye  row  0.59  mm  long,  procurved; 
MOQ  length  0.26  mm,  front  width  0.22 
mm,    back    width   0.32   mm;    e\e    interdis- 


240         Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  4 


tances  (mm):  AME-AME  0.04,  AME- 
ALE  0.03,  PME-PME  0.14,  PME-PLE 
0.08,  ALE-PLE  0.04. 

Sternum  1.12  mm  long,  0.85  mm  wide. 
Chelicerae  0.65  mm  long  with  3  promar- 
ginal  teeth  and  6  retromarginal  denticles. 

Abdomen  2.11  mm  long,  1.31  mm  wide. 
Epigasti'ic  furrow  0.67  mm  from  tracheal 
spiracle,  spiracle  0.76  mm  from  base  of 
spinnerets. 

All  coxae  with  round  knobs  ventrally. 
Femora  II  and  III  with  patches  of  short, 
thick  setae  ventrally.  Tibial  lengths  (mm) 
and  indices:  I  2.00,  9;  II  1.69,  13;  III  1.30, 
17;  IV  1.87,  12.  Ventral  spination:  tibiae 
I  2-2-0,  II  1-2-0,  III,  IV  1-2-2;  metatarsi 
I,  II  2-2-0,  III,  IV  2-2-2. 

Palpus  as  in  Figures  65,  69. 

Female  (San  Diego  Co.,  Cahfornia). 
Coloration  as  in  male. 

Total  length  5.04  mm.  Carapace  2.25  mm 
long,  1.76  mm  wide,  cephalic  width  0.95 
mm,  clypeus  height  0.12  mm.  Eyes:  diam- 
eters (mm):  AME  0.10,  ALE  0.13,  PME 
0.10,  PLE  0.13;  anterior  eye  row  0.51  mm 
long,  straight;  posterior  eye  row  0.68  mm 
long,  procurved;  MOQ  length  0.28  mm, 
front  width  0.26  mm,  back  width  0.36  mm; 
eye  interdistances  (mm):  AME-AME 
0.06,  AME-ALE  0.03,  PME-PME  0.16, 
PME-PLE  0.08,  ALE-PLE  0.05. 

Sternum  1.28  mm  long,  0.90  mm  wide. 
Chelicerae  0.70  mm  long  with  teeth  as  in 
male. 

Abdomen  3.10  mm  long,  2.02  mm  wide. 
Epigastric  furrow  0.77  mm  from  tracheal 
spiracle,  spiracle  1.03  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  1.62,  14;  II  1.49,  15;  III  1.17, 
21;  IV  1.69,  14.  Ventral  spination  as  in 
male  save  tibiae  III  1-1-0. 

Epigynum  as  in  Figure  72,  internal  geni- 
talia as  in  Figure  76. 

Natural  history.  Mature  males  have  been 
taken  from  late  April  through  late  June, 
mature  females  from  mid-May  through 
late  September.  Nothing  is  known  of  the 
habits  of  this  species. 


Distribution.  Southern  Utah  south  to 
southern  California  and  Arizona  (Map  2). 

Anyphaena  accentuata  Group 

Diagnosis.  Members  of  this  group  can 
be  immediately  differentiated  from  the 
other  nearctic  Anyphaena  by  the  presence 
of  only  one  pair  of  ventral  spines  on  meta- 
tarsi I  and  II.  Only  one  species  occurs  in 
America  north  of  Mexico. 

Description.  Total  length  4-6  mm.  Cara- 
pace longer  than  wide,  narrowed  in  front 
to  less  than  half  its  maximum  width  in 
males,  to  slightly  more  than  half  in  females. 
Clypeus  height  roughly  equal  to  anterior 
median  eye  diameter.  Median  eyes  smaller 
than  laterals.  Procurved  posterior  eye  row 
longer  than  recurved  anterior  row.  Me- 
dian ocular  quadrangle  longer  than  wide 
in  front,  wider  in  back  than  long.  Anterior 
median  eyes  separated  by  less  than  their 
diameter,  closer  to  anterior  laterals.  Pos- 
terior medians  separated  by  1.5  times  their 
diameter,  closer  to  posterior  laterals.  An- 
terior laterals  separated  by  their  radius 
from  posterior  laterals.  Sternum  longer 
than  wide,  unmodified.  Chelicerae  with  3 
promarginal  teeth  and  5-7  retrolateral  den- 
ticles. Abdomen  longer  than  wide,  tracheal 
spiracle  midway  between  epigastric  furrow 
and  base  of  spinnerets.  Leg  formula  1423, 
legs  unmodified.  Metatarsi  I  and  II  with 
one  pair  of  ventral  spines.  Palpus  with 
short  median  apophysis,  short  conductor 
and  conspicuous  embolus.  Cymbial  groove 
compressed  to  retrolateral  side  of  cymbium. 
Epigynum  with  hood.  Internal  genitalia 
with  anterior  membranous  dorsal  cover. 

Variation.  No  significant  variation  was 
detected  in  Anyphaena  aperta. 

Anyphaena  accentuata  (Walckenaer) 
Figure  134 

Aranea  accentuata  Walckenaer,  1802,  Faun.  Paris, 
2:  226.    Type  lost,  presumed  destroyed. 

Anyphaena  accentuata,  Roewer,  1954,  Katalog  der 
Araneae,  2:  522.  Bonnet,  1955,  Bibliographia 
Araneoriini,  2:  338. 


Spider  Family  Axypiiaenidak  •  Plafnick        241 


A  drawing  of  tlic  palpus  of  tliis  Enro- 
pc^an  spider,  type  species  ol  tlie  genus  A/ij/- 
pJiaena,  is  included  for  pinposes  of  com- 
parison to  A.  aperta.  Confusion  exists 
between  AnypJiaena  accentuatii,  A.  ohscura 
(Sundex'all)  and  A.  sabina  L.  Koch,  and 
the  female  is  therefore  not  illustrated  and 
no  description  is  gi\^en.  The  male  illus- 
trated is  from  England. 

Anyphaena  aperta  (Banks) 
Map  2;  Figures  135-137 

Gaijenna  aperta  Banks,  1921,  Pioc.  California 
Acad.  Sci.,  11:  100,  fig.  ,3  (  9  ).  Female  holo- 
t\'pe  from  OKinpia,  Washington,  in  MCZ,  ex- 
amined. 

Ani/pliacna  aperta,  Bryant,  1931,  Psyche,  38:  114, 
pi.  8,  fig.  35,  9  .  Fox,  1938,  Iowa  State  Coll.  J. 
Sci.,  12:  238,  pi.  1,  fig.  6,  $.  Roewer,  1954, 
Katalog  der  Araneae  2:  528.  Bonnet,  1955, 
Bibliographia   Araneornm,   2:    342. 

Didiinosis.  In  addition  to  the  diagnostic 
character  of  the  species  group,  Amjphaemi 
aperta  can  readily  be  distinguished  from 
all  other  North  American  anyphaenids  by 
the  sharply  pointed  median  apophysis  of 
males  (Fig.  135)  and  the  membranous  dor- 
sal cover  of  the  internal  genitalia  of  females 
(Fig.  137).  Although  the  distribution  indi- 
cates that  this  might  be  an  inti-oduced 
species,  no  specimens  or  described  species 
from  the  Palearctic  or  Oriental  regions  re- 
semble Anyphaena  aperta. 

Male  (Yamhill  Co.,  Oregon).  Total 
length  4.32  mm.  Carapace  1.98  mm  long, 
1.63  mm  wide;  cephalic  width  0.74  mm, 
clypeus  height  O.OS  mm,  light  orange- 
brown,  darker  towards  sides,  with  two 
dark  paramedian  longitudinal  bands.  Eyes: 
diameters  (mm):  AME  0.07,  ALE  0.11, 
PME  0.09,  PLE  0.11;  anterior  eye  row 
0.44  mm  long,  recurved;  posterior  eye  row 
0.62  mm  long,  procurved;  MOQ  length 
0.26  mm,  front  width  0.20  mm,  back  width 
0.32  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.03,  PME-PME 
0.14,  PME-PLE  0.10,  ALE-PLE  0.06. 

Sternum  1.04  mm  long,  0.89  mm  wide, 
pale  orange   \\'ith    translucent   border   and 


darkened  extensions  to  coxae.  Chelicerae 
0.55  mm  long  with  3  promarginal  teeth  and 
5  retromarginal  denticles,  orange-brown 
with  boss  outlined  in  gray.  Labium  and 
endites  pale  orange,  darkest  proximally. 
Endites  not  invaginated. 

Abdomen  2.52  mm  long,  1.51  mm  wide, 
pale  white  with  transx'crse  rows  of  dark 
markings,  venter  pale  with  a  clump  of 
thick  elongate  setae  posteriorly.  Epigastric 
furrow  0.86  mm  from  tracheal  spiracle, 
spiracle  0.74  mm  from  base  of  spinnerets. 

Legs  pale  yellow,  unmodified.  Tibial 
lengths  (mm)  and  indices:  I  1.87,  12;  II 
1.70,  13;  III  1.27,  18;  IV  1.73,  14.  Ventral 
spination:  tibiae  I,  II  2-2-2,  III  1-2-2, 
IV  2-2-2;  metatarsi  I,  II  2-0-0,  III  2-0-2, 
IV  2-2-2. 

Palpus  as  in  Figure  135. 

Female  (Curry  Co.,  Oregon).  Colora- 
tion as  in  male. 

Total  length  5.83  mm.  Carapace  2.65 
mm  long,  2.05  mm  wide,  cephalic  width 
1.17  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.10,  ALE  0.12, 
PME  0.12,  PLE  0.13;  anterior  eye  row 
0.61  mm  long,  slightly  recurved;  posterior 
eye  row  0.87  mm  long,  procurved;  MOQ 
length  0.35  mm,  front  width  0.30  mm,  back 
width  0.44  mm;  eye  interdistances  (mm): 
AME-AME  0.09,  AME-ALE  0.05,  PME- 
PME  0.19,  PME-PLE  0.14,  ALE-PLE  0.07. 

Sternum  1.46  mm  long,  1.04  mm  wide. 
Chelicerae  0.80  mm  long  with  3  promar- 
ginal teeth  and  7  retromarginal  denticles. 

Abdomen  4.00  mm  long,  2.60  mm  wide, 
without  thick  setae  ventrally.  Epigastric 
furrow  0.81  mm  from  tracheal  spiracle, 
spiracle  1.03  mm  from  base  of  spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  1.87, 
16;  II  1.77,  16;  III  1.31,  22;  IV  1.87,  17. 
Ventral  spination  as  in  male  except  tibiae 
I  2-2-0  and  IV  1-2-2. 

Epigynum  as  in  Figure  136,  internal 
genitalia  as  in  Figure  137. 

Natural  history.  Mature  males  have  been 
taken  from  late  Marcli  through  early  Sep- 
tember,  mature   females   from   ],\tc   March 


242         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


through  early  November.  Specimens  have 
been  taken  from  redwoods  and  red  cedars. 
Distribution.  Pacific  coast  from  British 
Coknnbia  south  to  southern  Cahfornia 
(Map  2). 

Wulfila  O.  P.-Cambridge 

WuIfiJa  O.  P.-Cambridge,  1895,  Biologia  Central! 
Americana,  Aran.,  1:  158.  Type  species  Wulfila 
pallidtis  O.  P.-Cambridge,  1895,  designated  by 
Simon,  1897,  Hist.  Natur.  Araign.,  2:   103. 

Cragits  O.  P.-Cambridge,  1896,  Biologia  Centrali 
Americana,  Aran.,  1:  215.  Type  species  by  mono- 
typy  Cragiis  pallidus  O.  P.-Cambridge,  1896. 
NEW  SYNONYMY. 

Anyphaenella  Bryant,  1931,  Psyche,  38:  115.  Type 
species  by  original  designation  Clubiona  salta- 
hunda  Hentz,    1847.     NEW   SYNONYMY. 

Diagnosis.  Wulfila  may  be  easily  recog- 
nized by  their  long,  thin,  pale  white  legs. 
Leg  I  in  particular  is  greatly  elongated, 
with  its  tibial  index  usually  5  or  less.  Pal- 
pal structure  indicates  that  this  genus  is 
closely  related  to  Amjphaena.  There  are 
probably  more  than  fifty  species  in  this 
genus;  most  occur  in  Central  America  and 
the  West  Indies. 

Description.  Total  length  2.5-4.5  mm. 
Carapace  longer  than  wide,  narrowed  in 
front  to  from  one-half  to  two-thirds  its 
maximum  width.  Clypeus  height  greater 
than  anterior  median  eye  diameter.  Pos- 
terior median,  posterior  lateral  and  anterior 
lateral  eyes  subequal  in  size,  somewhat 
larger  than  anterior  medians.  Procurved 
posterior  eye  row  longer  than  straight  an- 
terior row.  Median  ocular  quadrangle 
twice  as  wide  in  back  as  in  front.  Anterior 
median  eyes  separated  by  less  than  their 
diameter,  by  roughly  their  diameter  from 
anterior  laterals.  Posterior  medians  sepa- 
rated by  almost  twice  their  diameter,  by 
their  diameter  from  posterior  laterals.  An- 
terior laterals  separated  by  roughly  their 
diameter  from  posterior  laterals.  Sternum 
longer  than  wide,  unmodified.  Chelicerae 
with  3-6  promarginal  teeth,  often  on  ca- 
rina, and  5-10  retromarginal  denticles. 
Abdomen  longer  than  wide,  tracheal  spira- 


cle midway  between  epigastric  furrow  and 
base  of  spinnerets.  Leg  formula  1423,  legs 
long,  thin,  pale  white.  Leg  I  greatly  elon- 
gated. Metatarsi  I  and  II  with  two  pairs 
of  ventral  spines.  Coxae  of  males  often 
with  spurs  and  knobs;  leg  III  spination 
often  reduced.  Palpus  with  an  elongated 
median  apophysis,  enlarged  conductor  and 
conspicuous  embolus.  Retrolateral  tibial 
apophysis  greatly  expanded  except  in  W. 
wunda.  Epigyna  and  internal  genitalia 
small  and  diverse. 

Variation.  None  of  the  species  in  this 
genus  show  any  significant  individual  or 
geographic  intraspecific  variation  in  struc- 
ture, size  or  coloration. 

Key  to  Species 

la.  Carapace   and   abdomen   with    dark   mark- 
ings    saltabunda 

lb.  Carapace     and     abdomen     without     dark 
markings  2 

2a.  Males     3 

2b.  Females . 7 

3a.  At  least  one  pair  of  coxae  modified  with 
spurs  or  knobs  4 

3b.  All  coxae  unmodified  alba 

4a.  Coxae  I  and/or  II  modified  with  spurs  or 
knobs     5 

4b.  Coxae  III  and/or  IV  modified  with  spurs 
or  knobs   - 6 

5a.  Retrolateral    tibial    apophysis    more    than 
half  the  tibial  length   (Fig.  93)  —  bryantae 

5b.   Retrolateral  tibial  apophysis  less  than  half 
the  tibial  length  ( Fig.  95 )   ._  wunda 

6a.   Retrolateral    tibial    apophysis    greatly    ex- 
panded at  tip   (Fig.  86)   ..._.  tantilh 

6b.   Retrolateral    tibial    apophysis    not    greatly 
expanded  at  tip  (Fig.  88)  immaculella 

7a.  Epigynum  with  long  ducts   (Figs.  91,  97, 
98 )    8 

7b.  Epigynum  without  long  ducts    (Figs.   90, 
96)    : 10 

8a.  Epigynum     with     a    heart-shaped     atrium 
(Fig.   97)    wunda 

8b.  Epigynum  without  a  heart-shaped  atrium     9 

9a.  Epigynal  ducts  terminating  far  anterior  of 
epigynal  openings   ( Fig.  91 )   tantilla 

9b.  Epigynal   ducts   temiinating  near  epigynal 

openings  ( Fig.  98 )  immaculella 

10a.  Epigynum    with    anterolateral   flaps,    with- 
out a  medial  ridge   (Fig.  90)   alba 

10b.  Epigynum  without  anterolateral  flaps,  with 
a  medial  ridge  (Fig.  96)  bryantae 


Spideh  Family  Anyphaenidaf,  •  Platnick        243 


^; 

V) 

i 
) 

!-•- 

\U^ 

•^ 

•, 

rf  ? 

• 

^  _-  •^-^•.^% 

•\ 

\/^ 

Wulfila  saltabundQ^ 

1 

--—— ^^"^ 

^^-j 

^ 

\ 

Ifila  tantilla 

1 

V 

i 

r 

i 

1 

1 

■^ 

Wulfila  immaculella    | 

VV 

\ 

U  '^ 


— u-^/-     ... 


r 


Wulfllo  albo 


\--  - 

1    '        ■ 

1             —  -'i 

V     .' 

N 

. 

^"^     /^ 

v-4 

/■•f 

\ 

\ 

Wulfllo  bryantae 

"\ 

Map  3.     Distributions  of  Teudis  calcar,  Wulfila  alba,  W.  bryantae,  W.  immaculella,  W.  saltabunda,  W.  tantilla  and 
IV.  wunda. 


Wulfila  pallidas  O.  P.-Cambridge 
Figure  144 

Wulfila  palUdus  O.  P.-Cambridge,  1895,  Biologia 
Central!  Americana,  Aran.,  1:  159,  pi.  19,  fig. 
11  (  9  ).  Female  holotype  from  Teapa,  Ta- 
basco, Mexico,  in  BMNH,  examined.  Bonnet, 
1959,   Bibliographia  Araneorum,  2:   4832. 

Wulfila  pallida,  Simon,  1897,  Hist.  Natur.  Araign., 
2:  94.  Roewer,  1954,  Katalog  der  Araneae,  2: 
554. 

Vulfila  pallida,  Simon,  1897,  Hist.  Natur.  Araign., 
2:  103. 

Thi.s  Mexican  .species,  though  belonging 
to  a  distinct  species  group,  closely  resem- 
bles the  North  American  Wulfila  in  body 
form,  leg  length  and  coloration.  It  is  the 
type  .species  of  Wulfila. 

Wulfila  saltabunda  (Hentz), 
new  combination 
Map  3;  Figures  81,  82,  89,  99 

Cluhiona  saltabunda  Hentz,  1847,  J.  Boston  Soc. 
Natur.  Hist.,  5:  453,  pi.  23,  fig.  23  (  9  ).  Fe- 
male holotype  from  Alabama  in  Boston  Soc. 
Natur.  Hist.  (Boston  Mu.seum  of  Science),  de- 
stroyed by  beetles. 

Amjphaena  saltabunda,  Fmerton,  1890,  Trans. 
Connecticut  Acad.   Sci.,   8:    187,  figs.  4-4d,    $, 


$ .  Emerton,  1902,  Common  Spiders,  p.  14, 
figs.  46,  47,  5  ,  9  . 

Gayenna  saltabunda,  Comstock,  1912,  Spider 
Book,  p.  563,  figs.  638,  639,   $,   9 . 

Anyphaenella  saltabunda,  Bryant,  1931,  Psyche, 
38:  116,  pi.  7,  figs.  18,  22,  $,  9.  Comstock, 
1940,  Spider  Book,  rev.  ed.,  p.  576,  figs.  638, 
639,  $,  9.  Ka.ston,  1948,  Bull.  Connecticut 
Geol.  Natur.  Hist.  Surv.,  70:  406,  figs.  1465- 
1470,  $,  9.  Roewer,  1954,  Katalog  der 
Araneae,  2:  530.  Bonnet,  1955,  Bibliographia 
Araneorum,  2:  349. 

Dia<i,nosis.  Wulfila  saltabunda  is  the 
only  species  in  this  area  which  has  dark 
markings  on  the  carapace  and  abdomen. 
In  addition,  the  shape  of  the  retrolateral 
tibial  apophysis  ( Fig.  82 )  and  sperma- 
thecae  (Fig.  99)  serve  to  distinguish  it 
from  W.  alba,  its  closest  relative. 

Male  (Suffolk  Co.,  New  York).  Total 
length  3.06  mm.  Carapace  1.46  mm  long, 
1.04  mm  wide,  cephalic  width  0.54  mm, 
clypeus  height  0.07  mm,  pale  white  with 
thin  dark  border  and  two  dark  paramedian 
longitudinal  bands.  Eyes:  diameters 
(mm):  AME  0.05,  ALE  0.09,  PME  0.09, 
PLE  0.09;  anterior  eye  row  0.39  mm  long, 
straight;  posterior  eye  row  0.50  mm  long, 


244         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


86 


^■'^., 


■■    90 


Plate  7 


Figures  81,  83,  85,  87.  Left  palpi,  ventral  view.  Figures  82,  84,  86,  88.  Left  palpi,  retrolateral  view.  Figures 
89-91.  Epigyna,  ventral  view.  81.  82,  89.  Wulfila  saltabunda  (Hentz).  83,  84,  90.  Wulfila  alba  (Hentz).  85,  86, 
91.    Wulfila  tantilla  Chickering.     87,  88.    Wulfila  immaculella  (Gertsch). 


Spider  Faaiily  Anyphaenidae  •  Phifnick 


245 


procurx'ed;  MOQ  length  0.22  mm,  front 
width  0.14  mm,  back  width  0.30  mm;  eye 
interdistances  (mm):  AME-AME  0.04, 
AME-ALE  0.03,  PME-PME  0.12,  PiME- 
PLE  0.06,  ALE-PLE  0.03. 

Sternum  0.86  mm  long,  0.59  mm  wide, 
pale  white  with  thick  tran.slucent  border 
witli  extensions  to  coxae  and  large  triangu- 
lar dark  spots  between  coxae.  Chelicerae 
0.40  mm  long  with  6  promarginal  teeth  and 
7  retromarginal  denticles,  pale  white  with 
boss  outlined  in  gray  and  several  very  long 
setae.  Laliium  and  endites  pale  white, 
endites  not  in\'aginated  at  middle. 

Abdomen  1.60  mm  long,  0.97  mm  wide, 
pale  white  with  transverse  rows  of  dark 
spots,  venter  with  thin  dark  median  line 
anterior  of  epigastric  furrow  and  two  large 
median  dark  spots  between  epigastric  fur- 
row and  spinnerets.  Epigastric  furrow  0.40 
mm  from  tracheal  spiracle,  spiracle  0.45 
nmi  from  base  of  spinnerets. 

Legs  pale  white,  unmodified,  though  leg 
III  spination  reduced.  Tibial  length  (mm) 
and  indices:  I  2.70,  4;  II  1.42,  9;  III  1.08, 
13;  IV  1.55,  10.  Ventral  spination:  tibiae 
I  2-2-0,  II  1-1-0,  III  0-1-0,  IV  1-1-0; 
metatarsi  I,  II  2-2-0,  III  0-0-0,  IV  1-2-2. 

Palpus  as  in  Figures  81,  82. 

Female  (Suffolk  Co.,  New  York).  Col- 
oration as  in  male. 

Total  length  4.18  mm.  Carapace  1.78 
mm  long,  1.28  mm  wide,  cephalic  width 
0.70  mm,  clypeus  height  0.09  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.09, 
PME  0.09,  PLE  0.10;  anterior  eye  row  0.43 
mm  long,  straight;  posterior  eye  row  0.58 
mm  long,  procurved.  MOQ  length  0.28 
mm,  front  width  0.15  mm,  back  width  0.32 
mm;  eye  interdistances  (mm):  AME- 
AME  0.04,  AME-ALE  0.05,  PME-PME 
0.14,  PME-PLE  0.08,  ALE-PLE  0.04. 

Sternum  0.99  mm  long,  0.74  mm  wide. 
Chelicerae  0.56  mm  long  with  6  promar- 
ginal teeth  and  10  retromarginal  denticles. 

Abdomen  2.47  mm  long,  2.27  mm  wide. 
Epigastric  furrow  0.85  mm  from  tracheal 
spiracle,  .spiracle  0.85  mm  from  base  of 
spinnerets. 


Tibial  lengths  (mm)  and  indices:  I  2.56, 
5;  II  1.31,  12;  III  0.90,  20;  IV  1.67,  12. 
\'entral  spination  as  in  male  except  tibiae 
II  2-2-0  and  III  1-1-0  and  metatarsi  III 
2-1-0. 

Epigynum  as  in  Figure  89,  internal  geni- 
talia as  in  Figure  99. 

Natural  Jiistonj.  Mature  males  have  been 
taken  from  mid-April  through  late  August, 
mature  females  from  late  April  through 
late  August.  Specimens  have  been  taken 
by  sweeping  and  on  apple  trees. 

Distrilnifion.  Nova  Scotia  west  to  Min- 
nesota and  Nebraska,  south  to  Florida  and 
eastern  Texas  ( Map  3 ) . 

Wulfila  alba  (Hentz), 
new  combination 
Map  3;  Figures  83,  84,  90,  100 

Cluhiona    albeus    Hentz,     1847,     J.     Boston     See. 

Natur.  Hist.,  5:  454,  pi.  23,  fig.  24   {$).    Male 

holotype   from   Alabama  in   Boston   Soc.   Natur. 

Hist.     (Boston  Museum  of  Science),  destroyed 

by  beetles. 
Anyphaetia  alhcns,  Marx,  1883,  in  Howard,  A  List 

of  the  Invertebrate  Fauna  of  South  Carolina,  p. 

24. 
Chimcanthium  alhens,  Mar.x,  1890,  Proc.  U.S.  Nat. 

Mus.,  12:  513. 
AmiphaeucUa    alba,    Bryant,     1931,    Psyche,    38: 

116,  pi.  7,  figs.  20,  21,    S,    9.    Roewer,   1954, 

Katalog    der   Araneae,    2:    530.     Bonnet,    1955, 

Bibliographia  Araneorum  2:   349. 

Diagnosis.  Wulfila  alba  is  closest  to  W. 
sahaJninda  but  may  be  distinguished  from 
it  by  its  lack  of  dark  markings,  the  .spur- 
like retrolateral  tibial  apophysis  (Fig.  84) 
and  the  shape  of  the  spermathecae  (Fig. 
100). 

Male  (Orange  Co.,  Florida).  Coloration 
as  in  Wulfila  saltahunda  except  that  dark 
markings  are  entirely  absent. 

Total  length  3.65  mm.  Carapace  1.57 
mm  long,  1.21  mm  wide,  cephalic  width 
0.59  mm,  clypeus  height  0.07  mm.  Eyes: 
diameters  (mm):  AME  0.04,  ALE  0.07, 
PME  0.07,  PLE  0.07;  anterior  eye  row  0.36 
mm  long,  straight;  posterior  eye  row  0.49 
mm  long,  procurved;  MOQ  length  0.22 
mm,    front    width    0.12    mm,    back    width 


246         Bulletin  Museum  of  Comparative  Zoology,  Vol.  146,  No.  4 


0.26  mm;  eye  interdistances  (mm):  AME- 
AME  0.04,  AME-ALE  0.05,  PME-PME 
0.11,  PiME-PLE  0.06,  ALE-PLE  0.04. 

Sternum  0.95  mm  long,  0.70  mm  wide. 
Chelicerae  0.45  mm  long  with  6  promar- 
ginal  teeth  and  7  retromarginal  denticles. 

Abdomen  2.12  mm  long,  1.15  mm  wide. 
Epigastric  furrow  0.67  mm  from  tracheal 
spiracle,  spiracle  0.79  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  3.13, 
4;  II  1.85,  7;  III  1.12,  13;  IV  2.03,  6.  Ven- 
tral spination:  tibiae  I  2-2-0,  II  1-2-0,  III 
0-1-0,  IV  1-1-0;  metatarsi  I,  II,  III  2-2-0, 
IV  1-1-2. 

Palpus  as  in  Figures  83,  84. 

Female  (Indian  River  Co.,  Florida). 
Coloration  as  in  male. 

Total  length  4.00  mm.  Carapace  1.62 
mm  long,  1.28  mm  wide;  cephalic  width 
0.58  mm,  clypeus  height  0.06  mm.  Eyes: 
diameters  (mm):  AME  0.04,  ALE  0.07, 
PME  0.08,  PLE  0.08;  anterior  eye  row  0.40 
mm  long,  straight;  posterior  eye  row  0.54 
mm  long,  procurved;  MOQ  length  0.25  mm, 
front  width  0.14  mm,  back  width  0.29  mm; 
eye  interdistances  (mm):  AME-AME 
0.05,  AME-ALE  0.06,  PME-PME  0.13, 
PME-PLE  0.10,  ALE-PLE  0.06. 

Sternum  0.92  mm  long,  0.74  mm  wide. 
Chelicerae  0.41  mm  long  with  6  promar- 
ginal  teeth  and  9  retromarginal  denticles. 

Abdomen  2.66  mm  long,  1.51  mm  wide. 
Epigastric  furrow  0.68  mm  from  tracheal 
spiracle,  spiracle  0.88  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  3.13, 
4;  II  1.91,  8;  III  1.05,  13;  IV  2.07,  8.  Ven- 
tral spination  as  in  male  except  tibiae  II 
2-2-0  and  III  1-2-0  and  metatarsi  IV  2- 
2-2. 


Epigynum  as  in  Figure  90,  internal  geni- 
talia as  in  Figure  100. 

Natural  history.  Mature  males  have  been 
taken  from  late  March  through  early  Au- 
gust, mature  females  from  early  April 
through  late  August.  Specimens  have  been 
taken  by  sweeping,  on  pines,  and  in  Ma- 
laise and  pitfall  traps. 

Distribution.  Maryland  west  to  southern 
Illinois,  south  to  Florida  and  eastern  Texas 
(Map3). 

Wulfila  tantilla  Chickering 
Map  3;  Figures  85,  86,  91,  101 

Cragus  palUdus  O.  P. -Cambridge,  1896,  Biologia 
Centrali  Americana,  Aran.,  1:  215,  pi.  26,  fig. 
10  {  $).  Male  holotype  from  Santa  Ana,  Guate- 
mala, in  BMNH,  examined.  Preoccupied  by 
Wulfila  pallidus  O.  P. -Cambridge,  1895.  Roewer, 
1954,  Katalog  der  Araneae,  2:  535.  Bonnet, 
1956,   Bibliographia  Araneorum,   2:    1246. 

Wulfila  tantilla  Chickering,  1940,  Trans.  Amer. 
Microsc.  Soc,  59:  119,  figs.  64-66  {$).  Male 
holotype  from  El  Valle,  Panama,  in  MCZ, 
examined.  Roewer,  1954,  Katalog  der  Araneae, 
2:  555.   NEW  SYNONYMY. 

Wulfila  tenella  Chickering,  1940,  Trans.  Amer. 
Microsc.  Soc,  59:  120,  figs.  67,  68  (  $  ).  Fe- 
male holotype  from  El  Valle,  Panama,  in  MCZ, 
examined.  Roewer,  1954,  Katalog  der  Araneae, 
2:  555.   NEW  SYNONYMY. 

Diagnosis.  Wulfila  tantilla  is  very  closely 
related  to  W.  irnmaculella  but  may  be  dis- 
tinguished by  the  greath'  expanded  tip  of 
the  retrolateral  tibial  apophysis  (Fig.  86) 
and  by  the  epigynal  ducts  terminating  far 
anterior  of  the  epigynal  openings  (Fig. 
91). 

Male  (Webb  Co.,  Texas):  Coloration  as 
in  Wulfila  alba,  except  that  the  posterior 
declivity  of  the  carapace  is  darkened. 

Total  length  3.02  mm.  Carapace  1.62 
mm  long,  1.12  mm  wide,  cephalic  width 
0.63  mm,  clypeus  height  0.09  mm.    Eyes: 


Plate  8 
Figures  92,  94,  107.  Left  palpi,  ventral  view.  Figures  93,  95,  105.  Left  palpi,  retrolateral  view.  Figures  96-98, 
106.  Epigyna,  ventral  view.  Figures  99-104,  108.  Internal  genitalia,  dorsal  view.  Figure  109.  Body,  dorsal  view. 
92,  93,  96,  102.  Wulfila  bryantae  new  species.  94,  95,  97,  104.  Wulfila  wunda  new  species.  98,  103.  Wulfila 
irnmaculella  (Gertsch).  99.  Wulfila  saltabunda  (Hentz).  100.  Wulfila  alba  (Hentz).  101.  Wulfila  tantilla  Chick- 
ering.   105-109.    Oxysoma  cubana  Banl<s.    (Figs.  105,  106,  109  by  Wilton  Ivie,  not  to  scale.) 


Spider  Family  Anyphaenidae  •  Plalnick        247 


248         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


diameters  (mm):  AME  0.06,  ALE  0.07, 
PME  0.09,  PLE  0.09;  anterior  eye  row 
0.41  mm  long,  straight;  posterior  eye  row 
0.53  mm  long,  procurved;  MOQ  length 
0.23  mm,  front  width  0.16  mm,  back  width 
0.30  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.04,  PME-PME 
0.13,  PME-PLE  0.06,  ALE-PLE  0.04. 

Sternum  0.74  mm  long,  0.63  mm  wide. 
Chelicerae  0.58  mm  long  with  5  promar- 
ginal  teeth  on  a  carina  and  8  retromarginal 
denticles. 

Abdomen  1.57  mm  long,  0.97  mm  wide. 
Epigastric  furrow  0.50  mm  from  tracheal 
spiracle,  spiracle  0.44  mm  from  base  of 
spinnerets. 

Coxae  III  and  IV  with  two  small  knobs. 
Tibial  lengths  (mm)  and  indices:  I  2.77,  3; 
II  1.51,  9;  III  0.86,  15;  IV  1.51,  9.  Ventral 
spination:  tibiae  I,  II  2-2-0,  III,  IV  1-2-0; 
metatarsi  I,  II  2-2-0,  III,  IV  2-2-2. 

Palpus  as  in  Figures  85,  86. 

Female  (Hidalgo  Co.,  Texas).  Colora- 
tion as  in  male  of  Wulfila  alha. 

Total  length  2.92  mm.  Carapace  1.34 
mm  long,  0.99  mm  wide,  cephalic  width 
0.67  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.06, 
PME  0.06,  PLE  0.06;  anterior  eye  row  0.37 
mm  long,  straight;  posterior  eye  row  0.50 
mm  long,  procurved;  MOQ  length  0.20 
mm,  front  width  0.16  mm,  back  width  0.26 
mm;  eye  interdistances  ( mm ) :  AME- 
AME  0.05,  AME-ALE  0.04,  PME-PME 
0.13,  PME-PLE  0.06,  ALE-PLE  0.05. 

Sternum  0.89  mm  long,  0.61  mm  wide. 
Chelicerae  0.50  mm  long  with  4  promar- 
ginal  teeth  and  6  retromarginal  denticles. 

Abdomen  1.62  mm  long,  1.15  mm  wide. 
Epigastric  furrow  0.59  mm  from  tracheal 
spiracle,  spiracle  0.52  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  2.36,  5;  II  1.21,  11;  III  0.77, 
18;  IV  1.40,  11.  Ventral  spination  as  in 
male  save  tibiae  IV  1-1-0. 

Epigynum  as  in  Figure  91,  internal  geni- 
talia as  in  Figure  101. 

Natural  history.   Mature  males  have  been 


taken  from  mid-April  through  mid-Octo- 
ber, mature  females  apparently  year-round. 
Nothing  is  known  of  the  habits  of  this 
species. 

Distribution.  Southern  Texas  south  to 
the  Canal  Zone  (Map  3). 

Wulfila  immaculella  (Gertsch), 
new  combination 
Map  3;  Figures  87,  88,  98,  103 

Amjphac'uella  immaculella  Gertsch,  1933,  Amer. 
Mus.  Novitates,  No.  637:  9,  fig.  14  (  ?  ).  Fe- 
male holotype  from  Sabino  Basin,  Santa  Cata- 
lina  Movmtains,  Arizona,  in  AMNH,  examined. 
Roewer,  1954,  Katalog  der  Araneae,  2:  530. 
Bonnet,  1955,  Bibliographia  Araneorum,  2:  349. 

Diagnosis.  Wulfila  immaculella  is  very 
closely  related  to  W.  tantilla  but  may  be 
distinguished  by  the  unexpanded  tip  of  the 
retrolateral  tibial  apophysis  (Fig.  88)  and 
by  the  epigynal  ducts  terminating  near  the 
epigynal  openings  (Fig.  98). 

Male  (Sonora,  Mexico).  Coloration  as 
in  Wulfila  alba. 

Total  length  3.60  mm.  Carapace  1.64 
mm  long,  1.12  mm  wide,  cephalic  width 
0.67  mm,  clypeus  height  0.08  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.08, 
PME  0.08,  PLE  0.08;  anterior  eye  row  0.40 
mm  long,  straight;  posterior  eye  row  0.52 
mm  long,  procurved;  MOQ  length  0.22 
mm,  front  width  0.14  mm,  back  width 
0.31  mm;  eye  interdistances  (mm):  AME- 
AME  0.05,  AME-ALE  0.05,  PME-PME 
0.14,  PME-PLE  0.08,  ALE-PLE  0.04. 

Sternum  0.90  mm  long,  0.68  mm  wide. 
Chelicerae  0.51  mm  long  with  4  promar- 
ginal  teeth  and  5  retromarginal  denticles. 

Abdomen  2.07  mm  long,  1.00  mm  wide. 
Epigastric  furrow  0.63  mm  from  tracheal 
spiracle,  spiracle  0.74  mm  from  base  of 
spinnerets. 

Coxae  III  with  one,  coxae  IV  with  two 
small  knobs.  Tibial  lengths  (mm)  and  in- 
dices: I  3.42,  3;  II  2.05,  5;  III  1.30,  11;  IV 
2.11,  8.  Ventral  spination:  tibiae  I  2-2-0, 
II  1-2-0,  III  0-1-0,  IV  1-1-0;  metatarsi  I, 
II  2-2-0,  III,  IV  2-1-2. 

Palpus  as  in  Figures  87,  88. 


Spider  Family  Anyphaenidae  •  Plat  nick        249 


Fenmle  (Sonora,  Mexico).  Coloration  as 
in  male  of  Wiilfila  alba. 

Total  length  3.64  mm.  Carapace  1.5S 
mm  long,  1.08  mm  wide,  cephalic  width 
0.61  mm,  clypen.s  height  0.10  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.07, 
PME  0.08,  PLE  0.07;  anterior  eye  row 
0.40  mm  long,  straight;  posterior  eye  row 
0.53  mm  long,  procurved;  MOQ  length 
0.22  mm,  front  width  0.15  mm,  back  width 
0.30  mm;  eye  interdistances  (mm):  AME- 
AiME  0.05,  AME-ALE  0.04,  PME-PME 
0.12,  PME-PLE  0.08,  ALE-PLE  0.04. 

Sternum  0.94  mm  long,  0.71  mm  wide. 
Chelicerae  0.53  mm  long  with  teeth  as  in 
male. 

Abdomen  2.05  mm  long,  1.40  mm  wide. 
Epigastric  furrow  0.65  mm  from  tracheal 
spiracle,  spiracle  0.74  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  2.81,  4;  II  1.62,  9;  III  0.92, 
15;  IV  1.75,  8.  Ventral  spination  as  in  male 
except  tibiae  II  2-2-0  and  III  1-1-0. 

Epigynum  as  in  Figure  98,  internal  geni- 
talia as  in  Figure  103. 

Natural  Jiistory.  Mature  males  have  been 
taken  in  July,  mature  females  in  June  and 
July.    One  male  was  taken  on  Platanus  sp. 

Distrihuiion.  Southern  Arizona  and  So- 
nora (Map  3). 

Wulfila  bryantae  new  species 
Map  3;  Figures  92,  93,  96,  102 

Types.  Male  holotype,  female  paratype 
from  5  miles  east  of  Edinburg,  Hidalgo 
Co.,  Texas,  20  April  1937  (S.  Mulaik),  de- 
posited in  AMNH.  Male  and  female  para- 
types  from  Jim  Wells  and  Cameron  Coun- 
ties, Texas,  deposited  in  MCZ.  The 
specific  name  is  a  patronym  in  honor  of 
Miss  Elizabeth  Bryant,  in  recognition  of  her 
pioneering  work  on  North  American  any- 
phaenids. 

Diagnosis.  Wulfila  bryantae  is  a  distinc- 
tive species  easily  recognized  by  its  stubby 
median  apophysis  (Fig.  92)  and  the  medial 
ridge  on  the  epigynum   (Fig.  96). 


Male  (Hidalgo  Co.,  Texas).  Coloration 
as  in  Wulfila  alba. 

Total  length  3.35  mm.  (Carapace  1.44 
mm  long,  1.08  mm  wide,  cephalic  width 
0.81  mm,  clypeus  height  0.10  mm.  Eyes: 
diameters  (mm):  AME  0.06,  ALE  0.06, 
PME  0.07,  PLE  0.07;  ant(>rior  eye  row  0.49 
mm  long,  slightly  rcx-urved;  posterior  eye 
row  0.62  mm  long,  procurved;  MOQ  length 
0.26  mm,  front  width  0.20  mm,  back  width 
0.30  mm;  eye  interdistances  (mm):  AME- 
AME  0.07,^  AME-ALE  0.07,  PME-PME 
0.15,  PME-PLE  0.13,  ALE-PME  0.06. 

Sternum  0.97  mm  long,  0.55  mm  wide. 
Chelicerae  0.73  mm  long  with  3  promar- 
ginal  teeth  on  a  carina  and  7  retromarginal 
denticles. 

Abdomen  1.80  mm  long,  1.12  mm  wide. 
Epigastric  fvnrow  0.56  mm  from  tracheal 
spiracle,  spiracle  0.68  mm  from  base  of 
spinnerets. 

Coxae  I  with  a  small  knob,  coxae  II  with 
two  spurs.  Tibial  lengths  (mm)  and  in- 
dices: I  2.76,  4;  II  1.85,  7;  III  0.92,  15;  IV 
1.89,  7.  \'entral  spination:  tibiae  I,  II  2- 
2-0,  III  1-2-0,  IV  1-1-0;  metatarsi  I,  II  2- 
2-0,  III,  IV  2-1-2. 

Palpus  as  in  Figures  92,  93. 

Female  (Hidalgo  Co.,  Texas).  Colora- 
tion as  in  male  of  Wulfila  alba. 

Total  length  3.78  mm.  Carapace  1.44 
mm  long,  0.99  mm  wide,  cephalic  width 
0.74  mm,  clypeus  height  0.07  mm.  Eves: 
diameters  (mm):  AME  0.06,  ALE  6.07, 
PME  0.06,  PLE  0.07;  anterior  eye  row 
0.42  mm  long,  slightly  recurved;  posterior 
eye  row  0.59  mm  long,  procurved;  MOQ 
length  0.23  mm,  front  width  0.17  mm,  back 
width  0.27  mm;  eye  interdistances  (mm): 
AME-AME  0.06,  AME-ALE  0.05,  PME- 
PME  0.14,  PME-PLE  0.13,  ALE-PLE  0.06. 

Sternum  0.74  mm  long,  0.64  mm  wide. 
Chelicerae  0.62  mm  long  with  5  promar- 
ginal  teeth  and  5  retromarginal  denticles. 

Abdomen  2.59  mm  long,  2.16  nun  wide. 
Epigastric  furrow  0.90  mm  from  tracheal 
spiracle,  spiracle  0.88  mm  from  base  of 
spinnerets. 


250         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  2.24,  5;  II  1.37,  9;  III  0.72, 
19;  IV  1.35,  10.  Ventral  spination  as  in 
male  except  tibiae  III,  IV  2-2-0  and  meta- 
tarsi III,  IV  2-0-2. 

Epigynum  as  in  Figure  96,  internal 
genitalia  as  in  Figure  102. 

Natural  history.  Mature  males  have  been 
taken  from  late  April  through  early  June, 
mature  females  from  early  April  through 
early  December.  Nothing  is  known  of  the 
habits  of  this  species. 

Distribution.  Southern  Texas  and  Ta- 
maulipas  (Map  3). 

Wulfila  wunda  new  species 
Map  3;  Figures  94,  95,  97,  104 

Wulfila  immaculata,  Bryant  (not  Banks),  1936, 
Psyche,  43:  98,  fig.  1,  $.  Male  allotype  from 
Brichell  Hammock,  Florida  Keys,  in  MCZ, 
examined.  Not  Wulfila  immaculata  Banks,  1914, 
Bull.  Amer.  Mus.  Natur.  Hist.,  33:  640,  pi.  43, 
fig.  7,  9  .  Female  holotype  from  Vinales,  Pinar 
del  Rio,  Cuba,  in  AMNH,  examined. 

Types.  Male  holotype,  female  paratype 
from  Tavernier,  Monroe  Co.,  Florida,  16 
February  1951  (A.  M.  Nadler),  deposited 
in  AMNH.  Male  and  female  paratypes 
from  Dade  Co.,  Florida,  deposited  in  MCZ. 
The  specific  name  is  an  arbitrary  combina- 
tion of  letters. 

Diagnosis.  Wulfila  wunda  is  a  distinc- 
tive species  the  genitalia  of  which  are 
quite  different  from  those  of  the  other 
Wulfila  in  America  north  of  Mexico:  the 
retrolateral  tibial  apophysis  is  very  short 
( Fig.  95 )  and  the  epigynum  has  an  atrium 
(Fig.  97). 

Male  (Dade  Co.,  Florida).  Coloration 
as  in  Wulfila  alba. 

Total  length  3.42  mm.  Carapace  1.55 
mm  long,  1.08  mm  wide,  cephalic  width 
0.68  mm,  clypeus  height  0.06  mm.  Eyes: 
diameters  (mm):  AME  0.05,  ALE  0.07, 
PME  0.08,  PLE  0.08;  anterior  eye  row  0.48 
mm  long,  straight;  posterior  eye  row  0.59 
mm  long,  procurved;  MOQ  length  0.20 
mm,  front  width  0.14  mm,  back  width  0.29 
mm;    eye    interdistances     (mm):      AME- 


AME  0.05,  AME-ALE  0.09,  PME-PME 
0.14,  PME-PLE  0.12,  ALE-PLE  0.04. 

Sternum  1.06  mm  long,  0.70  mm  wide. 
Chelicerae  0.85  mm  long  with  4  promar- 
ginal  teeth  and  6  retromarginal  denticles. 

Abdomen  1.91  mm  long,  1.01  mm  wide. 
Epigastric  furrow  0.70  mm  from  ti'acheal 
spiracle,  spiracle  0.76  mm  from  base  of 
spinnerets. 

Coxae  II  with  a  small  knob.  Tibial 
lengths  (mm)  and  indices:  I  4.10,  3;  II 
1.87,  8;  III  1.01,  15;  IV  2.05,  7.  Ventral 
spination:  tibiae  I,  II  2-2-0,  III  0-1-0,  IV 
0-2-0;  metatarsi  I,  II  2-2-0,  III  0-2-0,  IV 
2-1-2. 

Palpus  as  in  Figures  94,  95. 

Female  (Dade  Co.,  Florida).  Colora- 
tion as  in  male  of  Wulfila  alba. 

Total  length  3.74  mm.  Carapace  1.55 
mm  long,  1.15  mm  wide,  cephalic  width 
0.72  mm,  clypeus  height  0.07  mm.  Eyes: 
diameters  (mm):  AME  0.04,  ALE  0.06, 
PME  0.07,  PLE  0.07;  anterior  eye  row  0.49 
mm  long,  straight;  posterior  eye  row  0.59 
mm  long,  procurved;  MOQ  length  0.20 
mm,  front  width  0.15  mm,  back  width 
0.30  mm;  eye  interdistances  ( mm ) :  AME- 
AME  0.06,  AME-ALE  0.10,  PME-PME 
0.16,  PME-PLE  0.12,  ALE-PME  0.04. 

Sternum  0.90  mm  long,  0.67  mm  wide. 
Chelicerae  0.65  mm  long  with  5  promar- 
ginal  teeth  and  9  retromarginal  denticles. 

Abdomen  2.16  mm  long,  1.15  mm  wide. 
Epigastric  furrow  0.74  mm  from  tracheal 
spiracle,  spiracle  0.83  mm  from  base  of 
spinnerets. 

Legs  unmodified.  Tibial  lengths  (mm) 
and  indices:  I  3.13,  4;  II  1.44,  10;  III  0.76, 
20;  IV  1.58,  9.  Ventral  spination  as  in  male 
except  tibiae  III  1-2-0  and  IV  0-1-0  and 
metatarsi  III  1-2-0  and  IV  1-2-2. 

Epigynum  as  in  Figure  97,  internal  geni- 
talia as  in  Figure  104. 

Natural  history.  Mature  males  have  been 
taken  from  mid-February  through  mid- 
May,  mature  females  apparently  year- 
round.  Nothing  is  known  of  the  habits  of 
this  .species. 


Spider  Family  Anyphaenidae  •  Platnick        251 


Distrihuiion.  Southern  Florida,  Culxi, 
and  Mona  Island  (  Map  3). 

Aysha  Keyserling 

Aysha  Keyserling,  1891,  Spinn.  Ainer.  ( Brasil. 
Spiiin.),  3:  83,  129.  Type  species  Aysha  pros- 
pera  Keyserling,  1891,  designated  by  Simon, 
1897,  Hist.  Natm-.  Araig.,  2:  104. 

Diagnosis.  Aysha  is  easily  recognized  by 
the  greatly  adxanced  placement  of  the 
tracheal  spiracle,  located  just  behind  the 
epigastric  furrow.  The  genitalic  structure 
is  quite  different  from  that  of  Amjphacna 
and  Wulfila  and  the  genus  undoubtedly 
represents  a  different  evolutionary  line. 
There  are  probably  more  than  thirty  spe- 
cies in  this  genus;  they  occur  commonly 
in  both  North  and  South  America. 

Description.  Total  length  4-9  mm.  Cara- 
pace longer  than  wide,  narrowed  in  front 
to  more  than  half  its  maximum  width. 
Clypeus  height  roughly  equal  to  anterior 
median  eye  diameter.  All  eyes  subequal  in 
size.  Procurved  posterior  eye  row  longer 
than  recurved  anterior  row.  Median  ocular 
quadrangle  longer  than  wide  in  front, 
wider  in  back  than  long.  Anterior  median 
eyes  separated  by  slightly  less  than  their 
diameter,  slightly  closer  to  anterior  laterals. 
Posterior  medians  separated  by  up  to  twice 
their  diameter,  closer  to  posterior  laterals. 
Anterior  laterals  separated  by  their  radius 
from  posterior  laterals.  Sternum  longer 
than  wide,  unmodified.  Chelicerae  with  3- 
4  promarginal  teeth  and  7-9  retromarginal 
denticles.  Abdomen  longer  than  wide, 
tracheal  spiracle  much  closer  to  epigastric 
furrow  than  to  base  of  spinnerets.  Leg 
formula  1423,  legs  unmodified.  Metatarsi  I 
and  II  with  one  pair  of  \'entral  spines.  Pal- 
pus with  greatly  enlarged  base  of  embolus, 
long  curving  embolus  and  short  conductor. 
Ventral  tibial  apophysis  sometimes  present 
in  addition  to  retrolateral  tibial  apophysis. 
Epigynum  with  anterior  median  opening 
and  two  sidepieces.  Internal  genitalia  with 
long,  sometimes  coiling,  ducts. 

Variation.     Only    Aysha    gracilis    shows 


significant  variation,  and  that  is  in  size  and 
not  strnetine  or  coloration.  The  size  of  both 
the  whole  animal  and  ol  the  genitalia  vary 
geographically.  The  largest  .specimens 
( males  with  cymbium  length  averaging  1.3 
mm)  occur  in  Virginia  and  surrounding 
states,  with  smaller  individuals  occurring 
in  the  north  (New  England  and  Michigan 
males  with  cymbium  length  averaging  1.1 
mm)  and  in  the  south  (Texas  males  with 
cymbium  length  averaging  0.9  mm). 

Key  to  Species 

la.  Males    2 

lb.  Females    -..- 7 

2a.  Palpus  without  a  ventral  tibial  apophysis 
(VTA)    (Figs.  Ill,  119)  -.._ 3 

2b.  Palpus  with  a  ventral  tibial  apophysis 
(VTA),  sometimes  small,  transparent,  eas- 
ily overlooked  (Figs.  113,  115,  117,  121)     4 

3a.  Embolus  restricted  to  distal  half  of  palpal 
bulb    (Fig.    118)    arunda 

3b.  Embolus  not  restricted  to  distal  half  of  pal- 
pal bulb    (Fig.    110)    velox 

4a.  VTA  erect,  sclerotized,  relatively  large 
(Figs.  113,  115)  5 

4b.  VTA  recumbent,  transparent,  relativelv 
small  (Figs.  117,  121)  '.  6 

5a.  Distal  retrolateral  tip  of  tegulum  with  a 
flap  covering  embolus   (Fig.   112)        decepta 

5b.  Distal  retrolateral  tip  of  tegulum  with  a 
sharp  point  underlying  embolus  (Fig.  114) 
— incursa 

6a.  Base  of  embolus  recurved,  with  a  sharp 
spike    (Fig.    120)    camhridgei 

6b.  Base  of  embolus  not  recurved,  forming  a 
smooth   arc    (Fig.    116)    fitacilis 

7a.  Internal  genitalia  with  simple  uncoiled 
ducts   (Figs.   124,   127,  141,   143)   9 

7b.  Internal  genitalia  coiled  or  with  accessory 
ducts  (Figs.  125,  142) 8 

8a.  Internal  genitalia  highly  coiled  ( Fig. 
125 )    '...... vclox 

8b.  Internal  genitalia  not  coiled  but  with  loop- 
ing accessory  ducts   (Fig.   142)   arunda 

9a.  Median  epig\'nal  opening  near  anterior 
rim  (Figs.  123,  126,  138)  10 

91).   Median   epigynal   opening  near  middle   of 

epigynum    ( Fig.   140 )    -..  gracilis 

10a.   Median  epig\nal  opening  much  wider  than 

epigynal  sidepieces   (Fig.   138)       cand)ridgc'i 
lOb.   Median   epigynal   opening   not   wider  than 

epigynal  sidepieces   (Figs.   123,   126)    11 

11a.  Base  of  epig>nal  sidepieces  near  epigastric 
furrow  (Fig.  126);  internal  genitalia  with 
angular  ducts    (Fig.    127)    incursa 


252         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


.  ^       Aysha  gracilis 


I / 


;      1     /         HM 


~T— 7 
I 


Ayshc  velox 


r? 


i? 


♦^^'^r 


cQ^* 


i 

i 

"  1              \ 

r' 

) 

( 

-A 

Aysha  arunda 

.„         V 

Map  4.     Distributions  of  Aysha  arunda.  A.  cambridgei,  A.    decepta,  A.  gracilis,  A.  incursa  and  A.  velox. 


111).  Base  of  epigynal  sidepieces  far  from 
epigastric  furrow  (Fig.  123);  internal 
genitalia  with  rounded  ducts  (Fig.  124) 
decepta 

Aysha  prospers  Keyserling 
Figure  145 

Ay.sha  prospera  Keyserling,  1891,  Spinnen  Amer- 
ikas  (Brasil.  Spinn.),  3:  129,  pi.  4,  fig.  88 
(  9  ).  Female  holotype  from  Rio  Grande,  Brasil, 
in  BMNH,  examined.  Roewer,  1954,  Katalog 
der  Araneae,  2:  533.  Bonnet,  1955,  Biblio- 
graphia    Araneorum,    2:     838. 

This  South  American  species,  type  spe- 
cies of  Aysha,  is  a  member  of  a  large,  dis- 
tinct species  group.  Somatic  characters 
clearly  ally  it  with  the  North  American 
forms  included  in  the  genus. 

Aysha  gracilis  (Hentz) 

Map  4;  Figures  116,  117,  140,  143 

Chihiona  gracilis  Hentz,  1847,  J.  Boston  Soc. 
Natur.  Hist.,  5:  452,  pi.  23,  fig.  9(5).    Type 


specimens  from  North  Carolina  and  Alabama  in 
Boston  Soc.  Natur.  Hist.  (Boston  Museum  of 
Science),  destroyed  by  beetles. 

Anijphaena  gracilis,  L.  Koch,  1836,  Arach.  Fam. 
brassidae,  p.  195,  pi.  8,  fig.  130,  9  .  Comstock, 
1912,  Spider  Book,  p.  561,  fig.  633,  $  (not 
fig.  632). 

Anijphaena  rubra  Emerton,  1890,  Trans.  Connecti- 
cut Acad.  Sci.,  8:  186,  pi.  6,  fig.  1(9).  Male 
allotype  (?)  from  Franklin  Park,  Boston,  Mas- 
sachusetts, in  MCZ,  examined.  Emerton,  1909, 
Trans.  Connecticut  Acad.  Sci.,  14:  220,  pi.  9, 
fig.  8-8c,  $ . 

Aysha  gracilis,  Bryant,  1931,  Psyche,  38:  119,  pi. 
7,  fig.  13,  pi.  8,  fig.  26,  $,  9.  Chickering, 
1939,  Pap.  Michigan  Acad.  Sci.,  24:  53,  figs. 
9-11,  $,  9.  Comstock,  1940,  Spider  Book, 
rev.  ed.,  p.  575,  fig.  633,  $  (not  fig.  632). 
Kaston,  1948,  Bull.  Connecticut  Geol.  Natur. 
Hist.  Surv.,  70:  405,  figs.  1452,  1459-1464,  $, 
9  .  Roewer,  1954,  Katalog  der  Araneae,  2:  534. 
Bonnet,  1955,  Bibliographia  Araneorum,  2:  837. 

Diagnosis.  Aysha  gracilis  is  closest  to 
A.  cambridgei  but  lacks  the  sharp  spike  on 
the  proximal  edge  of  the  base  of  embolus 
(Fig.   116)   of  that  .species.    Females  have 


Spider  Family  Anyphaenidae  •  Platnick        253 


Plate  9 
Figures  110,  112,   114,  116.    Left  palpi,  ventral  view.    Figures   111,   113,   115,   117 
110,   111.    Aysha   velox   (Becker).     112,   113.    Aysha   decepta  (Banks) 
116,  117.    Aysha  gracilis  (Hentz). 


Left  palpi,   retrolateral  view/. 
114,  115.    Aysha  incursa  (Chamberlin). 


the  median  epigynal  opening  near  the  mid-  long,   2.02  mm   wide,   cephalic  width   1.17 

die  of  the  epigynum  (Fig.  140).    Variation  mm,  clypeus  heigiit  0.09  mm,  light  orange- 

in  thi.s  species  is  discussed  above.  brown,  darkest  anteriorly,  with   thin   dark 

Male    (Middlesex    Co.,    Massachusetts),  border  and  two  dark  paramedian  longitudi- 

Total  length  5.73  mm.    Carapace  2.56  mm  nal  bands.    Eyes:    diameters  (mm):    AME 


254 


Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.  4 


0.09,  ALE  0.11,  PME  0.09,  PLE  0.11;  an- 
terior eye  row  0.60  mm  long,  slightly  re- 
cnrved;  posterior  eye  row  0.80  mm  long, 
procurved;  MOQ  length  0.32  mm,  front 
width  0.26  mm,  back  width  0.38  mm;  eye 
interdistances  (mm):  AME-AME  0.09, 
AME-ALE  0.07,  PME-PME  0.19,  PME- 
PLE  0.15,  ALE-PLE  0.06. 

Sternum  1.44  mm  long,  1.08  mm  wide, 
light  orange-brown  with  translucent  border 
and  darkened  extensions  to  coxae.  Chelic- 
erae  1.12  mm  long  with  4  promarginal 
teeth  and  8  retromarginal  denticles,  dark 
orange-brown  proximally,  dark  brown  dis- 
tally.  Labium  and  endites  light  orange- 
brown,  darkest  proximally.  Endites  sharply 
invaginated  at  middle. 

Abdomen  3.20  mm  long,  1.73  mm  wide, 
pale  grayish-brown  with  transverse  rows 
of  dark  markings,  venter  pale.  Epigastric 
furrow  0.40  mm  from  tracheal  spiracle, 
spiracle  1.73  mm  from  base  of  spinnerets. 

Legs  light  orange-brown  with  distal  seg- 
ments darkest.  Tibial  lengths  (mm)  and 
indices:  I  2.64,  10;  II  1.87,  15;  III  1.19,  26; 
IV  2.09,  15.  Vential  spination:  tibiae  I,  II 
2-2-2,  III  1-2-2;  IV  2-2-2;  metatarsi  I,  II 
2-0-0,  III  2-1-2,  IV  2-2-2. 

Palpus  as  in  Figure  116,  117. 

Female  (Washington  Co.,  Arkansas). 
Coloration  as  in  male. 

Total  length  8.42  mm.  Carapace  2.75 
mm  long,  2.11  mm  wide,  cephalic  width 
1.47  mm,  clypeus  height  0.10  mm.  Eyes: 
diameters  (mm):  AME  0.14,  ALE  0.14, 
PME  0.13,  PLE  0.14;  anterior  eye  row  0.43 
mm  long,  recurved;  posterior  eye  row  1.04 
mm  long,  procurved;  MOQ  length  0.43 
mm,  front  width  0.36  mm,  back  width  0.49 
mm;  eye  interdistances  (mm):  AME- 
AME  0.09,  AME-ALE  0.08,  PME-PME 
0.22,  PME-PLE  0.18,  ALE-PLE  0.06. 

Sternum  1.84  mm  long,  1.31  mm  wide. 
Chelicerae  1.57  mm  long  with  teeth  as  in 
male. 

Abdomen  5.76  mm  long,  3.53  mm  wide. 
Epigastric  furrow  0.68  mm  from  tracheal 
spiracle,  spiracle  3.24  mm  from  base  of 
spinnerets. 


Tibial  lengths  (mm)  and  indices:  I  2.56, 
16;  II  1.94,  20;  III  1.26,  30;  IV  2.30,  17. 
Ventral  spination:  tibiae  I  2-2-0,  II  1-2-1, 
III  1-1-2,  IV  1-2-2;  metatarsi  I,  II  2-0-0, 
III  2-0-2,  IV  2-2-2. 

Epigynum  as  in  Figure  140,  internal 
genitalia  as  in  Figure  143. 

Natural  history.  Mature  males  and  fe- 
males have  been  taken  year-round.  Speci- 
mens have  been  taken  by  sweeping,  in 
pitcher  plants,  on  loblolly  pine,  in  fall  web- 
worm  nests  and  frequently  in  houses. 

Distribution.  New  England  west  to  Wis- 
consin and  Iowa,  south  to  Florida  and  east- 
ern Texas  ( Map  4 ) . 

Aysha  cam  bridge!  Bryant 

Map  4;  Figures  120,  121,  138,  141 

Aysha  cambiidgei  Bryant,  1931,  Psyche,  38:  119, 
pi.  7,  fig.  15  {  $  ).  Male  holotype  from 
Guanajuato,  Mexico,  in  MCZ,  examined.  Roe- 
wer,  1954,  Katalog  der  Araneae,  2:  532.  Bon- 
net, 1955,  Bibliographia  Araneorum,  2:   836. 

Diagnosis.  Aysha  cambridgei  is  closely 
related  to  A.  gracilis  but  has  a  distinctive 
spike  on  the  proximal  edge  of  the  base  of 
the  embolus  (Fig.  120)  and  the  median 
epigynal  opening  near  the  anterior  rim  of 
the  epigynum  ( Fig.  138 ) . 

Male  (Jeff  Davis  Co.,  Texas).  Colora- 
tion as  in  Aysha  gracilis  except  that  the  ab- 
domen is  pale  white  with  two  dark  para- 
median longitudinal  bands. 

Total  length  5.87  mm.  Carapace  2.41 
mm  long,  1.91  mm  wide,  cephalic  width 
0.97  mm,  clypeus  height  0.11  mm.  Eyes: 
diameters  (mm):  AME  0.11,  ALE  0.12, 
PME  0.11,  PLE  0.11;  anterior  eye  row  0.57 
mm  long,  recurved;  posterior  eye  row  0.75 
mm  long,  procurved;  MOQ  length  0.33 
mm,  front  width  0.28  mm,  back  width  0.38 
mm;  eye  interdistances  (mm):  AME- 
AME  0.06,  AME-ALE  0.05,  PME-PME 
0.16,  PME-PLE  0.11,  ALE-PLE  0.05. 

Sternum  1.42  mm  long,  1.01  mm  wide. 
Chelicerae  0.98  mm  long  with  4  promar- 
ginal teeth  and  7  retromarginal  denticles. 

Abdomen  3.49  mm  long,  1.58  mm  wide. 
Epigastric  furrow  0.68  mm  from  tracheal 


Spideu  Family  Anyphaenidae  •  Platnick        255 


122 


126 


127 


125 

Plate  10 
Figures   118,   120.     Left  palpi,   ventral   view.     Figures   119,   121.     Left  palpi,   retrolateral   view.    Figures   122,   123, 
126.    Epigyna,  ventral  view.     Figures  124,  125,  127.    Internal  genitalia,  dorsal  view.     118,  119.    Aysha  arunda  new 
species.     120,   121.    Aysha   cambridgei   Bryant.      122,    125.    Aysha   velox  (Becker).     123,   124.    Aysha   decepta 
(Banks).     126,  127.   Aysha  incursa  (Chamberlin). 


.spiracle,    spiracle    1.55   mm    from    base    ot  Ventral   spination:     tibiae  I,   II  2-2-2,   III 

spinnerets.  1-2-2,  IV  2-2-2;  metatarsi  I,  II  2-0-0,  III, 

Tibial  lengths  (mm)  and  indices:    I  3.06,  IV  2-2-2. 

8;   II   1.87,   13;   III   1.28,  21;   IV  2.16,   12.  Palpus  as  in  Figures  120,  121. 


256         Bulletin  Museum  of  Comparative  Zoology,  Vol.   146,  No.   4 


Female  (Henderson  Co.,  Texas).  Color- 
ation as  in  male. 

Total  length  8.50  mm.  Carapace  3.35 
mm  long,  2.52  mm  wide,  cephalic  width 
1.69  mm,  clypeus  height  0.12  mm.  Eyes: 
diameters  (mm):  AME  0.14,  ALE  0.16, 
PME  0.14,  PLE  0.14;  anterior  eye  row  0.84 
mm  long,  recm'ved;  posterior  eye  row  1.11 
mm  long,  procurved;  MOQ  length  0.42 
mm,  front  width  0.37  mm,  back  width  0.50 
mm;  eye  interdistances  (mm):  AME- 
AME  0.10,  AME-ALE  0.07,  PME-PME 
0.22,  PME-PLE  0.20,  ALE-PLE  0.05. 

Sternum  1.91  mm  long,  1.22  mm  wide. 


'&' 


Chelicerae  1.69  mm  long  with  teeth  as  in 
male. 

Abdomen  5.04  mm  long,  2.88  mm  wide. 
Epigastric  furrow  0.61  mm  from  tracheal 
spiracle,  spiracle  3.17  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  12.88, 
12;  II  2.07,  17;  III  1.40,  26;  IV  2.57,  15. 
Ventral  spination  as  in  male  except  tibiae 
I,  II  2-2-0  and  III  2-2-2. 

Epigynum  as  in  Figure  138,  internal 
genitalia  as  in  Figure  141. 

Natural  history.  Mature  males  have  been 
taken  from  mid-June  through  early  August, 
mature  females  from  late  May  through 
early  August.  Specimens  have  been  taken 
on  trees  and  shrubs. 

Distribution.  South  central  states  from 
Alabama  to  western  Texas,  south  to  central 
Mexico  (Map  4). 

Aysha  decepta  (Banks) 

Map  4;  Figures  112,  113,  123,  124 

Amjphaena  decepta  Banks,  1899,  Proc.  Ent.  Soc. 
Washington,  4:  190.  Female  holotype  from 
Brazos  Co.,  Texas,  in  MCZ,  examined. 

Aysha  mimita  F.  O.  P.-Canibridge,  1900,  Biologia 
Centrali  Americana,  Aran.,  2:  99,  pi.  7,  figs. 
18-19  {  $,  ?  ).  Male  holotype,  female  allotype 
from  Guatemala,  in  BMNH,  examined.  Bryant, 
1931,  Psyche,  38:  120,  pi.  7,  fig.  17,  $.  Roe- 
wer,  1954,  Katalog  der  Araneae,  2:  533.  Bon- 
net, 1955,  Bibliographia  Araneorum,  2:  838. 
NEW  SYNONYMY. 

Aijsha  decepta,  Bryant,  1931,  Psyche,  38:  120,  pi. 
7,  fig.  16,  pi.  8,  fig.  27,    $,   9.    Roewer,  1954, 


Katalog    der   Araneae,    2:    534.     Bonnet,    1955, 
Bibliographia  Araneorum,  2:   836. 

Diagnosis.  Aysha  decepta  is  very  closely 
related  to  A.  incursa  but  has  a  characteris- 
tic flap  (on  the  retrolateral  tip  of  the  tegu- 
lum)  that  covers  the  embolus  (Fig.  112), 
while  the  base  of  the  epigynal  sidepieces 
is  a  considerable  distance  from  the  epigas- 
tric furrow  ( Fig.  123 ) .  Both  morphological 
and  zoogeographical  data  (Map  4)  indi- 
cate that  these  two  species  are  each  other's 
nearest  relatives. 

Male  (Hidalgo  Co.,  Texas).  Coloration 
as  in  Aysha  camhridgei. 

Total  length  4.82  mm.  Carapace  2.25 
mm  long,  1.76  mm  wide,  cephalic  width 
1.06  mm,  clypeus  height  0.10  mm.  Eyes: 
diameters  (mm):  AME  0.08,  ALE  0.10, 
PME  0.11,  PLE  0.11;  anterior  eye  row  0.58 
mm  long,  straight;  posterior  eye  row  0.75 
mm  long,  procurved;  MOQ  length  0.23 
mm,  front  width  0.24  mm,  back  width  0.39 
mm;  eye  interdistances  ( mm ) :  AME- 
AME  0.08,  AME-ALE  0.06,  PME-PME 
0.18,  PME-PLE  0.12,  ALE-PLE  0.05. 

Sternum  1.37  mm  long,  0.85  mm  wide. 
Chelicerae  0.97  mm  long  with  4  promar- 
ginal  teeth  and  7  retromarginal  denticles. 

Abdomen  2.74  mm  long,  1.39  mm  wide. 
Epigastric  furrow  0.38  mm  from  tracheal 
spiracle,  spiracle  1.28  mm  from  base  of 
spinnerets. 

Tibial  lengths  (mm)  and  indices:  I  2.54, 
9;  II  1.67,  14;  III  1.01,  25;  IV  1.89,  16. 
Venti-al  spination:  tibiae  I  2-2-0,  II  1-2-0, 
III  1-2-2,  IV  2-2-2;  m